Saddleback toad
The saddleback toads are a genus of tiny toads and frogs in the family Brachycephalidae in the order Anura, ranging from south Bahia to Santa Catarina 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, which once were placed in their own genus Psyllophryne. Some pumpkin toadlets are toxic and their often bright colours are considered aposematic. At about or less in snout–vent length, the flea frogs are some of the smallest frogs in the world.
The saddleback toads live among leaf litter in the Atlantic rainforest, ranging from near sea level to an altitude of, with most species restricted to highland cloud forest. They are mostly diurnal, feed on tiny invertebrates and breeding is by direct development, with eggs laid on land and hatching into young frogs.
The majority of the Brachycephalus species have only been discovered in the 21st century. Most species have tiny ranges and are seriously threatened.
Appearance
Morphology and size
The common name "saddleback toad" refers to the bony shields that cover their vertebrae, 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 morphology, and they mostly walk. The second group are the flea frogs or flea toads that have an appearance and morphology that is more like leptodactylid frogs, and they are better jumpers. The name "flea frog" is also sometimes used for the unrelated Adelophryne. The skin of the saddleback toads can be rugose 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.Brachycephalus are very small frogs and adults range from 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 in adults, but overlapping to some extent with the smallest species of pumpkin toadlets. Adults of the pumpkin toadlets are at least, with most species about, and the largest like B. darkside and B. margaritatus at least. At, B. pulex has the smallest minimum and maximum adult size reported in the genus. B. dacnis is similarly tiny, with a minimum snout–vent length of. This makes them the smallest known frogs, with only the northeast Brazilian Adelophryne michelin, and Cuban Eleutherodactylus iberia and E. limbatus approaching their tiny dimensions. In other continents, the only similar-sized or slightly smaller frogs are a few species in the genera Mini, Paedophryne and Stumpffia. Newly hatched saddleback toads are much smaller than the already tiny adults. For example, they are typically in B. ephippium, a species where adults are at least.
Colour and toxicity
The different Brachycephalus 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.The flea frogs or flea toads 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.
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 Brazilian gold frog. 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, B. brunneus and B. curupira, are overall brown, while a few others are essentially all-brown or greenish above, 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.
The bright yellow, orange or reddish colours of many species in this genus are generally considered aposematic, but toxicity has only been studied in a few species. The brightly coloured B. ephippium and B. pernix have tetrodotoxin and similar toxins in their skin and organs, whereas B. nodoterga, which has a much more subdued colouring but still with some yellow-orange, appears to be non-toxic. It is unknown how certain Brachycephalus species attain the strong neurotoxins, but they are possibly absorbed from the small invertebrates they eat, as known from some poison dart frogs and mantella frogs, or produced by bacteria inside their body.
At least two brightly coloured species, B. ephippium and B. pitanga, have bones that are fluorescent, which is visible through the skin of the toadlet when exposed to UV light. It was initially speculated that the fluorescent colour also is aposematic or that it is related to mate choice, 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.
Voice and hearing
Although not very loud for a frog/toad, Brachycephalus 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 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 territory from other males, but in most cases voice or visual displays are sufficient and fights are avoided.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 males when close. The calls of the different species differ to various extent in frequency, speed, structure and other features.
Peculiarly, at least the brightly coloured pumpkin toadlets B. ephippium and B. pitanga 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 vocal sac that inflates when calling, mouth gaping and waving of their arms. It is speculated that their calling is a vestigiality from the ancestral form of the genus, whereas their reduced hearing ability is a novel change in these species. Sounds make them more vulnerable to predators, but there has likely been little direct evolutionary pressure to lose it because of their toxicity. Unlike B. ephippium, B. pitanga 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 pumpkin toadlet B. actaeus and the dull-coloured flea frog B. hermogenesi strongly suggest that they are able to hear their own calls.
Behavior
Saddleback toads live in the leaf litter on forest floors, but on occasion may move to higher perches up to 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 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.Overall the saddleback toads are mostly active during the day, but some activity also occurs during the night and some species possibly are mostly nocturnal.
Reproduction
In general, saddleback toads are reported to use axillary or nearly axillary amplexus 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 B. ephippium will use two, beginning with an inguinal amplexus and only later changing to a nearly axillary position.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 on land. 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 tadpole or other aquatic stage.