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A new species of brachycephalid frog is described from the highlands of the Atlantic Forest in Serra do Mar, municipality of Cunha, São Paulo State, southeastern Brazil. Specimens were collected amidst the leaf litter at approximately 920 m above sea level. The new species is distinguished from all of its congeners by the combination of the following characters: snout–vent length 9.5–15.6 mm; rough dorsum; general background color orange with a variable density of whitish spots on dorsum; skin on head and dorsum granular with dermal ossification; skull, spinal processes of sacral and presacral vertebrae, and process of the fourth vertebra hyperossified; absence of hyperossification in the central portion of the skull; dorsal surface of body covered by osteoderms. The new species was found active by day and is locally abundant. Males hold territories during the reproductive season. Females had few mature oocytes during the same period. The advertisement call of B. crispus is a long and low-intensity buzz with a regular repetition of notes.
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... Several additional morphological features have been used for diagnosing species within the B. ephippium group, such as their dorsal coloration (e.g. Pombal & Gasparini, 2006;Alves et al., 2009;Haddad et al., 2010;Clemente-Carvalho et al., 2012;Condez et al., 2014;Nunes et al., 2021), osteological (Pombal, 2010;Campos et al., 2010;Pombal & Izecksohn, 2011;Condez et al., 2014;Guimarães et al., 2017;Nunes et al., 2021) and muscle pigmentation features (Guimarães et al., 2017;Nunes et al., 2021). However, recent molecular studies have demonstrated that the systematics of the group is divergent in few analysis, for example, in the phylogenetic position of B. hermogenesi and B. sulfuratus (Clemente-Carvalho et al., 2011;Padial et al., 2014;Firkowski et al., 2016;Monteiro et al., 2018;Condez et al., 2020;Dos Reis et al., 2020). ...
... Several additional morphological features have been used for diagnosing species within the B. ephippium group, such as their dorsal coloration (e.g. Pombal & Gasparini, 2006;Alves et al., 2009;Haddad et al., 2010;Clemente-Carvalho et al., 2012;Condez et al., 2014;Nunes et al., 2021), osteological (Pombal, 2010;Campos et al., 2010;Pombal & Izecksohn, 2011;Condez et al., 2014;Guimarães et al., 2017;Nunes et al., 2021) and muscle pigmentation features (Guimarães et al., 2017;Nunes et al., 2021). However, recent molecular studies have demonstrated that the systematics of the group is divergent in few analysis, for example, in the phylogenetic position of B. hermogenesi and B. sulfuratus (Clemente-Carvalho et al., 2011;Padial et al., 2014;Firkowski et al., 2016;Monteiro et al., 2018;Condez et al., 2020;Dos Reis et al., 2020). ...
... The presence or absence of bony plates is commonly reported in species descriptions and, although this feature has proven useful for species diagnosis, the morphology of these bony plates has not been investigated comparatively (but see Campos et al., 2010;Clemente-Carvalho et al., 2012;Dos Reis et al., 2020). Detailed osteological descriptions are known for 17 species (46%) of this genus (Ribeiro et al., 2005;Alves et al., 2006;Bornschein et al., 2016;Ribeiro et al., 2017;Monteiro et al., 2018;Nunes et al., 2021), with only eight species from the B. ephippium group (representing 22% of the genus and 61% of the group): B. bufonoides (Folly et al., 2020), B. crispus (Condez et al., 2014), B. darkside (Guimaraes et al., 2017), B. ephippium (Campos et al., 2010;Trewavas, 1932;Da Silva et al., 2007), B. guarani (Clemente-Carvalho et al., 2012), B. margaritatus (McLachlan, 1943; only skull features), B. toby (Haddad | 3 FOLLY et aL. et al., 2010) and Brachycephalus sp. ...
Article
Brachycephalus is a small, endemic genus of anurans that occur throughout the Brazilian Atlantic Forest. Recent analyses corroborated the monophyly of two species groups within this genus (B. ephippium and B. pernix), whereas the B. didactylus group appears to be polyphyletic. Herein, we compare and describe the skeletal system of all species from the Brachycephalus ephippium species group. We investigated diagnostic characters that are potentially useful to delimit similar species, confirmed the previously proposed genus definition and added six extra characters. We propose an osteological diagnosis for each species of the B. ephippium group, evaluating and describing intraspecific variations. Our results suggest that adults of B. ephippium group have ornamented parotic plates, ornamented post‐orbital crests, absence of quadratojugal and neopalatines, distal end of the otic ramus of squamosal expanded towards the parotic plate, reduced zygomatic ramus of the squamosals, posterolateral projection of the crista parotica, presence of ornamented spinal plates (except B. alipioi) and ornamented paravertebral plates. We highlighted the importance of including more than one specimen per species when dealing with miniaturized taxa. A comparison with other Brachycephalus species groups and with Ischnocnema was also provided. Lastly, we suggest some characters to be included in future phylogenetic analysis.
... mirissimus Pie, Ribeiro, Confetti, Nadaline, and Bornschein, 2018;Pie et al. 2018) lacked a comparison because the holotype B. atelopoide is lost; (2) another (B. pulex Napoli, Caramaschi, Cruz, and Dias, 2011;Napoli et al. 2011) was compared without citing the source of data; (3) Brachycephalus crispus Reis, 2014 (Condez et al. 2014) was compared using features that differ from those of the original description (Miranda-Ribeiro 1920) and redescription (Pombal 2010) of B. atelopoide; (4) the four remaining species descriptions included comparisons with the diagnosis of B. atelopoide. ...
... However, B. ephippium does not possess large and elongate warts. If B. atelopoide had an intermediate amo nt of perossification it would also resemble adult B. nodoterga, B. alipioi Pombal and Gasparini, 2006, B. crispus, B. guarani, B. pitanga Alves, Sawaya, Reis, and Haddad, 2009, B. toby Haddad, Alves, Clemente-Carvalho, and Reis, 2010, and B. vertebralis Pombal, 2001, all of which have lower perossification t an do ad t B. ephippium (Campos et al. 2010, Haddad et al. 2010, Clemente-Carvalho et al. 2012, Condez et al. 2014, 2020. In having large warts, B. atelopoide resembles B. nodoterga, B. pitanga, B. crispus (Campos et al. 2010, Condez et al. 2014, 2020, and especially B. pitanga, which has fewer of these warts. ...
... If B. atelopoide had an intermediate amo nt of perossification it would also resemble adult B. nodoterga, B. alipioi Pombal and Gasparini, 2006, B. crispus, B. guarani, B. pitanga Alves, Sawaya, Reis, and Haddad, 2009, B. toby Haddad, Alves, Clemente-Carvalho, and Reis, 2010, and B. vertebralis Pombal, 2001, all of which have lower perossification t an do ad t B. ephippium (Campos et al. 2010, Haddad et al. 2010, Clemente-Carvalho et al. 2012, Condez et al. 2014, 2020. In having large warts, B. atelopoide resembles B. nodoterga, B. pitanga, B. crispus (Campos et al. 2010, Condez et al. 2014, 2020, and especially B. pitanga, which has fewer of these warts. ...
... During the last ten years, researchers have looked deeper into these frogs' diversification and biogeography [6]. Consequently, the number of endemic Brachycephalus species increased significantly, especially with 15 species described for the last five years (e.g., B. crispus [7]; B. albolineatus [8]; B. sulfuratus [9]; B. darkside [10] do Meio Ambiente" (MMA), that also evaluates protocols for our collection and research. All collecting, holding and storage (scientific collection) procedures followed Brazilian animal care guidelines and were previously approved by the Universidade Estadual Paulista (UNESP) animal care committee (registration number CEUA IB/CLP #03/2020). ...
... The DF was acquired with "Peak frequency" parameter from Raven Pro 1.5 [41] and represents the frequency value that corresponds to the peak of energy within the call envelope. Our results were compared with call descriptions available for the Brachycephalus ephippium group: B. bufonoides (see [12]), B. crispus (see [7]), B. darkside (see [10]), and B. pitanga (see [42]). ...
... It is common to the majority of the species of Brachycephalus to present a restricted distribution due to the geographic pattern for the genus [5,80]. The sky islands scenario is supported by diversification-by-isolation and suggests that allopatric speciation due to climatic gradients is an important mechanism for generating species diversity and endemism in these regions [5,7,11,22]. However, not all of the species of this genus presents such restricted distributions, like B. nodoterga. ...
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Species of Brachycephalus has been having taxonomical issues due its morphological similarity and genetic conservatism. Herein, we describe a new species of Brachycephalus from the south Mantiqueira mountain range and semidecidual forests in the municipalities of Mogi das Cruzes, Campinas and Jundiaí, state of São Paulo, Brazil, based on an integrative approach. It can be distinguished from all species of the B. ephippium species group based on morphological characters (especially osteology and head shape), advertisement call and divergence in partial mitochondrial DNA gene sequences (16S). The new species is genetically similar to B. margaritatus and morphologically similar to B. ephippium. It can be differentiated from B. ephippium by the presence of dark faded spots on skull and post-cranial plates, presence of black connective tissue connective tissue scattered over dorsal muscu-lature, parotic plate morphology, smaller snout-vent length (adult SVL: males 13.46-15.92 mm; females 16.04-17.69 mm) and 3% genetic distance. We also present natural history data and discuss the robustness of the integrative approach, geographic distribution, genetic data, behaviour, fluorescence in ontogeny, and conservation status.
... All 14 described species of this group present dermal ossification, whereas three also present osteoderms (B. crispus Condez, Clemente-Carvalho et al., 2014, B. margaritatus Pombal Jr. & Izecksohn, 2011, B. nodoterga Miranda-Ribeiro, 1920 (Almeida-Silva et al., 2021;Campos et al., 2010;Clemente-Carvalho et al., 2009Condez et al., 2014Condez et al., , 2020Condez et al., 2021;Folly et al., 2020;Guimarães et al., 2017;Nunes et al., 2021;Pombal Jr. & Izecksohn, 2011). We argue that this speculation opens an avenue for future studies on whether these structures have a function analogous to the dorsal and lateral MDL expansions of certain species of Brachycephalus in protecting against desiccation. ...
... All 14 described species of this group present dermal ossification, whereas three also present osteoderms (B. crispus Condez, Clemente-Carvalho et al., 2014, B. margaritatus Pombal Jr. & Izecksohn, 2011, B. nodoterga Miranda-Ribeiro, 1920 (Almeida-Silva et al., 2021;Campos et al., 2010;Clemente-Carvalho et al., 2009Condez et al., 2014Condez et al., , 2020Condez et al., 2021;Folly et al., 2020;Guimarães et al., 2017;Nunes et al., 2021;Pombal Jr. & Izecksohn, 2011). We argue that this speculation opens an avenue for future studies on whether these structures have a function analogous to the dorsal and lateral MDL expansions of certain species of Brachycephalus in protecting against desiccation. ...
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The mineralized dermal layer (MDL) is found in most terrestrial anurans. Its thickness represents on average up to 8% of that of the entire skin. It has been proposed that it may reduce body water loss, act on homeostasis, support skin structure, or conversely, it may be a currently functionless trait constrained by groups' evolutionary history. We described the MDL morphology of 11 Brachy-cephalus species, terrestrial, miniaturized and microendemic anurans, and tested for its relationship with climate of higher latitude regions of the Atlantic Rainforest of Brazil. All species presented MDL, described with two distinct morphological patterns: homogeneous or heterogeneous MDL, the latter distinguishable by MDL with dorsal or lateral expansion, sometimes up to the limit of the epidermis and comprising up to 50% of the thicknesses of the entire skin. Climate differed between locations by MDL morphological group, less rainy or seasonally less rainy where species with heterogeneous MDL occur. Our results indicate that the abundance of calcium in MDL and its heterogeneous condition suggest its adaptive function in reduce water loss. Such adaptations in anurans in very humid highlands reinforce the mountains' propensity for rapid loss of humidity, demystifying them as an extremely abundant source of water. This is the third study that tested the relationship between the MDL morphology and the environment where species occur and the first that correlated this structure with the climate of anurans of the same habit and distributed in a single habitat, the Atlantic Rainforest.
... nov. overlaps those described for B. crispus and B. pitanga (Condez et al. 2014;Oliveira and Haddad 2017). ...
... Additionally, the transverse processes of vertebrae IV and V fused and associated to paravertebral plates in the new species is distinct from the condition of B. garbeanus, in which the transverse processes of vertebrae IV-VII are fused and associated to the larger paravertebral plates (see the specimen identified as B. ephippium from Nova Friburgo, RJ, in Campos et al. 2010). The absence of small dotted osteoderms distinguishes the new species from B. margar-itatus, in which the dorsal skin exhibits well-developed and sparse dotted osteoderms(Pombal and Izecksohn 2011;Condez et al. 2014). The discrete spinal plates along the main spinal axis distinguish the new species from B. darkside, in which the ornamentation on vertebrae VII and VIII are well developed and clearly visible externally(Guimarães et al. 2017). ...
Article
Four species of pumpkin toadlets are currently recognized by the extreme condition of hyperossification in the skull and vertebral column within the genus Brachycephalus. In addition to their larger body size, relative to other congeneric species, Brachycephalus darkside, Brachycephalus ephippium, Brachycephalus garbeanus, and Brachycephalus margaritatus share remarkable osteological features, such as the large paravertebral plates forming an ornamented dorsal bone shield, visible through the integument. We add to the current knowledge of this group by analyzing its diversity and describing a new hyperossified species from some important Atlantic Forest remnants in southeastern Brazil, including those bordering São Paulo, the largest and most populous city in South America. The new species is diagnosed by a combination of morphological, osteological, and bioacoustic characters, with further evidence of mtDNA sequences to confirm its distinction from the other congeners. The parotic and paravertebral bone plates externally bordered with a rough and pale contour can readily distinguish the new species from the other hyperossified species within Brachycephalus. The new species is widely distributed along the Serra do Mar mountain range, including the regionally named Serra de Paranapiacaba, in elevations from 700 to 1000 m above sea level (a.s.l.), in the central coast of the state of São Paulo, southeastern Brazil. We discuss valuable morphological characters within this group of species, reinforce the need for further studies, and highlight the relevance of protected areas for biodiversity conservation in metropolitan regions.
... Uncontrolled jumping with delayed hindlimb recovery and slow walking presumably place miniaturized frogs at greater risk of predation. Thus, we expect that miniaturization in frogs will often be accompanied by antipredator defense strategies such as toxicity, osteoderms, bony plates, aposematic coloration, or camouflage, all of which are seen in Brachycephalus (35,(40)(41)(42)(43). Given the overwhelming importance of scaling in determining fundamental aspects of organismal design, it is likely that future studies of locomotion in miniaturized frogs will reveal additional adaptations and tradeoffs that will further inform our understanding of life at the lower limits of vertebrate body size. ...
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Miniaturization has evolved repeatedly in frogs in the moist leaf litter environments of rainforests worldwide. Miniaturized frogs are among the world's smallest vertebrates and exhibit an array of enigmatic features. One area where miniaturization has predictable consequences is the vestibular system, which acts as a gyroscope, providing sensory information about movement and orientation. We investigated the vestibular system of pumpkin toadlets, Brachycephalus (Anura: Brachycephalidae), a clade of miniaturized frogs from Brazil. The semicircular canals of miniaturized frogs are the smallest recorded for adult vertebrates, resulting in low sensitivity to angular acceleration due to insufficient displacement of endolymph. This translates into a lack of postural control during jumping in Brachycephalus and represents a physical constraint resulting from Poiseuille's law, which governs movement of fluids within tubes.
... Uncontrolled jumping with delayed hindlimb recovery and slow walking presumably place miniaturized frogs at greater risk of predation. Thus, we expect that miniaturization in frogs will often be accompanied by antipredator defense strategies such as toxicity, osteoderms, bony plates, aposematic coloration, or camouflage, all of which are seen in Brachycephalus (35,(40)(41)(42)(43). Given the overwhelming importance of scaling in determining fundamental aspects of organismal design, it is likely that future studies of locomotion in miniaturized frogs will reveal additional adaptations and tradeoffs that will further inform our understanding of life at the lower limits of vertebrate body size. ...
Full-text available
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Miniaturization has evolved repeatedly in frogs in the moist leaf litter environments of rainforests worldwide. Miniaturized frogs are among the world's smallest vertebrates and exhibit an array of enigmatic features. One area where miniaturization has predictable consequences is the vestibular system, which acts as a gyroscope, providing sensory information about movement and orientation. We investigated the vestibular system of pumpkin toadlets, Brachycephalus (Anura: Brachycephalidae), a clade of miniaturized frogs from Brazil. The semicircular canals of miniaturized frogs are the smallest recorded for adult vertebrates, resulting in low sensitivity to angular acceleration due to insufficient displacement of endolymph. This translates into a lack of postural control during jumping in Brachycephalus and represents a physical constraint resulting from Poiseuille's law, which governs movement of fluids within tubes.
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Background: We are experiencing a global crisis in conservation, which has led to the prioritization of targets, such as nations, regions, and animal groups, which are necessary while resources are disputed. Brazil is a priority not only because of its megadiversity, high rates of endemism, and frequent descriptions of new species but also because of its high levels of deforestation. Among the species groups prioritized for conservation is the anurans (Amphibia: Anura), the population of which is severely declining. One group of anurans is the genus Brachycephalus, which includes 37 endemic species in the Brazilian Atlantic Rainforest. Some of these species have highly restricted distributions (<100 ha). Thirty new species have been described since 2000, and 55.3% of all species are threatened with extinction. Brachycephalus tridactylus was only recently described and remains restricted to its type locality. Because of its reduced geographical distribution (0.41 km2), it has been proposed to be considered as Vulnerable. The objective of this study is to reevaluate the conservation status of Brachycephalus tridactylus and propose conservation measures. Methods: We searched for new populations during 2016-2020, evaluated in loco impacts and potential impacts on the species' population, and performed an analysis of the density of this population and estimated its size. International Union for Conservation of Nature (IUCN) criteria were used to assess the conservation status of the species. Results: We recorded the species in seven new localities (from 715-1,140 m above sea level) in the state of São Paulo up to 33 km from the type locality of the species (in state of Paraná). We estimated the area of occupancy as 148.44 km2, densities as one calling male per 4.05 m2 and 130.00 m2, and a total population size of 4,429,722 adult individuals. Based on our finding, we proposed three lines of management: (1) formation of fire brigades, (2) management of residents' mules in the conservation unit and surrounding areas, and (3) management of degraded areas. We recommend changing the species' conservation status from Vulnerable to Endangered because of its fragmented distribution and decline in the area of occupancy and in the quality of its habitat. Our results have expanded the species previous geographic distribution and delimited areas without previous records. Our estimates of population density and size are in accordance with those verified for congeners. The conservation of this species benefits the environments and other species that inhabit them, being, therefore, strategic for receiving conservation actions that will spread throughout the ecosystem.
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The diminutive frogs of the genus Psyllophryne (Anura: Brachycephalidae) were known only from one species in the Atlantic Forest of the State of Rio de Janeiro, southeastern Brazil. Herein we describe a second species of Psyllophryne from coastal São Paulo State, southeastern Brazil. The new species can be distinguished from Psyllophryne didactyla mainly by the presence of a functional fifth toe. Pointed digit tips, vocal sac extending to midbelly, inferior margin of eye and anterior margin of nostril grooved, and reduction in size of fingers and number of toes are shared derived features of both Brachycephalus and Psyllophryne.
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A new species of Brachycephalus is described from Sacra Família do Tinguá, municipality of Engenheiro Paulo de Frontin, State of Rio de Janeiro, Atlantic Rain Forest. Brachycephalus margaritatus sp. nov. is characterized by large size (SVL 15.0 to 18.9 mm), orange color in life and cream to grayish cream in preservative, well developed dorsal plates with lateral edges curved down, and scattered bulges on upper surfaces of the body. The geographic distribution of B. margaritatus, B. ephippium, and B. garbeanus are provided.
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Dermal ossifications -- osteoderms -- are more common in anurans than has been generally acknowledged. In the hylid Phyllomedusa bicolor osteoderms are located in the dermis, cover the dorsal surface of the head and body, and are scattered on the lateral and ventral surfaces and the limbs. Each osteoderm consists of a vascularized bony basal plate (0.1 mm thick) from which bony lamellar spines protrude into the epidermis. The dorsal body osteoderms are approximately 3 mm2 in area. Similar osteoderms are present in Phyllomedusa vaillanti and Gastrotheca weilandii. In the pelobatid Megophrys nasuta osteoderms are present in the dorsal body skin and although comparable to the osteoderms of Phyllomedusa in size and shape they are histologically very different. Megophrys osteoderms are avascular and composed of calcified collagen bundles in an orderly three-dimensional arrangement. The leptodactylid Hylactophryne augusti also has small bony osteoderms in the skin of the dorsum. They differ from the hylid osteoderms and resemble Megophrys osteoderms in being avascular and having a matrix composed of horizontal and vertical bundles of collagen. Larger dermal bony dorsal plates are present in Lepidobatrachus and Ceratophrys (Leptodactylidae) and in Brachycephalus (Brachycephalidae). Anuran osteoderms are structurally different from, and not homologous with, caecilian dermal scales. Given the histological differences among the various anuran osteoderms and the taxonomic diversity (Hylidae, Pelobatidae, Leptodactylidae, and Brachycephalidae), osteoderms appear to have been independently evolved a number of times within the Anura. It is suggested that the term dermal scale be restricted to the bony scale of fishes and caecilians, and osteoderms be used to denote the dermal scutes of anurans and reptiles.