Next Article in Journal
Nuclear Orthologs Derived from Whole Genome Sequencing Indicate Cryptic Diversity in the Bemisia tabaci (Insecta: Aleyrodidae) Complex of Whiteflies
Next Article in Special Issue
Barcoding Analysis of Paraguayan Squamata
Previous Article in Journal
Genetic Data Suggest Multiple Introductions of the Lionfish (Pterois miles) into the Mediterranean Sea
Previous Article in Special Issue
Ecological and Conservation Correlates of Rarity in New World Pitvipers
 
 
Order Article Reprints
Font Type:
Arial Georgia Verdana
Font Size:
Aa Aa Aa
Line Spacing:
Column Width:
Background:
Article

Conservation Status of Brachycephalus Toadlets (Anura: Brachycephalidae) from the Brazilian Atlantic Rainforest

1
Instituto de Biociências, Universidade Estadual Paulista (UNESP), Praça Infante Dom Henrique s/no, São Vicente, São Paulo, CEP 11330-900, Brazil
2
Mater Natura—Instituto de Estudos Ambientais, Rua Lamenha Lins 1080, Curitiba, Paraná, CEP 80250-020, Brazil
3
Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Paraná, CEP 81531-980, Brazil
*
Author to whom correspondence should be addressed.
Diversity 2019, 11(9), 150; https://doi.org/10.3390/d11090150
Received: 1 July 2019 / Revised: 19 August 2019 / Accepted: 23 August 2019 / Published: 27 August 2019
(This article belongs to the Special Issue Systematics and Conservation of Neotropical Amphibians and Reptiles)

Abstract

:
The number of described anurans has increased continuously, with many newly described species determined to be at risk. Most of these new species inhabit hotspots and are under threat of habitat loss, such as Brachycephalus, a genus of small toadlets that inhabits the litter of the Brazilian Atlantic Rainforest. Of 36 known species, 22 were described in the last decade, but only 11 have been assessed according to the IUCN Red List categories, with just one currently listed as Critically Endangered. All available data on occurrence, distribution, density, and threats to Brachycephalus were reviewed. The species extent of occurrence was estimated using the Minimum Convex Polygon method for species with three or more records and by delimiting continuous areas within the altitudinal range of species with up to two records. These data were integrated to assess the conservation status according to the IUCN criteria. Six species have been evaluated as Critically Endangered, five as Endangered, 10 as Vulnerable, five as Least Concern, and 10 as Data Deficient. Deforestation was the most common threat to imperiled Brachycephalus species. The official recognition of these categories might be more readily adopted if the microendemic nature of their geographical distribution is taken into account.

1. Introduction

Frogs and toads (Anura) comprise more than 7000 species worldwide [1]. Special attention has been given to this group due to the large number of new species described each year as well as due to the increasing number of endangered species [2,3]. According to the IUCN Red List criteria [4,5], there are 1825 species of anurans at risk of extinction (25% of all species), making Anura the vertebrate order with the highest proportion of endangered species [5]. Since 1980, there have been records of a rapid population decline of nearly 450 anuran species [6,7,8]. The decline of these species can be mainly attributed to habitat loss and pathogens, such as chytrid fungi and Ranavirus [6,7,9,10,11]. Recently, Ranavirus has been reported in natural populations of frogs in South America, but the effects in wild anuran populations are still unknown [11]. Unlike Ranavirus, chytrid fungi (Batrachochytrium dendrobatidis) has been commonly reported as a cause of population decline in high altitude locations in Costa Rica and Panama [9]. Due to the rapid rate of the description of a new species, the proportion of endangered species, and sensitivity, Anura is the priority order for a conservation assessment, particularly in countries with a high level of deforestation, such as in Brazil [3].
The Atlantic Rainforest, a biodiversity hotspot [12], is the largest in area after the Amazon forest, with its original extent covering more than 1.3 million km2 [13,14]. It is located on the eastern coast of South America, stretching from northeastern to southern Brazil, with inland extensions to the east of Paraguay, northeast of Argentina, and central Brazil. This biome has been experiencing massive habitat loss due to agricultural expansion, urbanization, and historic loss of natural habitats [15]. Currently, only 28% of the original extent remains if secondary forests and forests affected by the edge effects are included [15]. The Atlantic Rainforest houses nearly 2500 species of vertebrates, including 550 anurans, of which 323 are endemic (63%) and 15 are currently considered to be threatened by extinction [1,5,16].
The genus Brachycephalus (Fitzinger, 1826) is endemic to the Atlantic Rainforest and includes small (less than 2.5 cm in snout-vent length) diurnal toadlets with a reduced number of digits, bright colors, neurotoxins in the skin, and direct development, and they live in leaf litter, specifically that of montane forests [17,18,19,20,21,22,23]. There are currently 36 recognized species of Brachycephalus [1], of which 22 have been described in the last decade [1]. Most have extremely restricted geographical distributions of less than 100 ha [12,24,25]. Brachycephalus is divided into three phenetic groups [26], two of which (B. ephippium and B. pernix groups) are montane with few records at lower altitudes, whereas the remaining group (B. didactylus group) includes more ecologically plastic species that occur from the sea level up to high altitudes [23,27]. The dependence on a colder climate and isolation in the mountains as sky islands have been hypothesized as the reason that montane groups have diverged into so many species (19 of B. pernix and 12 of B. ephippium groups), whereas the B. didactylus group includes only four species [23,28,29]. Another species (B. atelopoide) cannot be compared to any of the groups due to the unavailability of the holotype [23,30].
Species descriptions of Brachycephalus have not been accompanied by corresponding assessments of the conservation status. Only 11 species have been assessed for the IUCN Red List to date [31,32,33,34,35,36,37,38,39,40,41]: eight as Data Deficient (DD) and three as Least Concern (LC). The Ministério do Meio Ambiente (MMA, the Ministry of the Environment of the Brazilian government) evaluated only four species and categorized one as Critically Endangered (CR), two as DD, and one as Near Threatened (NT) [42,43,44,45]. The absence of conservation status assessments of most species and the evaluation of some of them as DD highlight the need for a comprehensive effort to assess the risk of extinction of the Brachycephalus species, most notably the microendemic taxa found in the B. pernix and B. ephippium species groups (sensu [26]). Species evaluated as DD should be prioritized to generate enough data to properly classify them into a conservation category [46,47].
One way to direct effective initiatives for conservation species is through a prior assessment of their conservation status [3]. There is a widely adopted IUCN methodology for proposing a conservation status [3], which serves an important role in allowing for comparisons and for classifying conservation actions as well the proposition of public policies. The objectives of the study were (1) to review data on occurrence, altitudinal distribution, density, and threats to the Brachycephalus species, (2) to compile new data from the literature and unpublished observations, (3) to generate systematized data on geographic distribution, population sizes, and threats to place them into IUCN conservation categories, and (4) to discuss conservation priorities and future management actions.

2. Material and Methods

All available occurrence records of Brachycephalus spp. were compiled from the literature up to the time of compilation (June 2019). The data encompassed toponymy, species identification, geographical coordinates of the occurrence record, and altitude of the corresponding site. Data on altitudinal range were also considered when available. The process began with the latest compilation of locality and altitude data for Brachycephalus provided by Bornschein et al. [23], and the same selection criteria were adopted for subsequent records. For example, those associated with precise localities were retained, and records that included only municipality names as occurrence information were discarded. Finally, the authors’ previously unpublished data were included.
Occurrence records were plotted using Google Earth Pro v. 7.1.4.1529 and connected to form a polygon using the Minimum Convex Polygon approach (MCP; [48]) with modifications suggested by Reinert et al. [49] and adopted by Bornschein et al. [23]. These modifications allow for the exclusion of inappropriate habitats, such as bodies of water, pastures, silvicultures, urban areas, rock areas, and/or forest areas, beyond the altitudinal range of occurrence of the species.
Polygon delimitation required three or more occurrence records. For species with one or two records, polygons encompassing the altitudinal range of the species were created [23]. A continuous topography inside the polygon was considered a location (sensu IUCN and as IUCN [48]) that could potentially contain one or more records of a given species. The topography was considered discontinuous if it was isolated by altitudes beyond the altitudinal range of the respective species.
The MCP and altitudinal polygons were measured using GEPath v. 1.4.5 to obtain the extent of occurrence (EO; IUCN [48]; see also [23,25,50]) of each species. Because some species have such reduced EO, they could potentially also be ranked by area of occupancy (AO), although AO was not measured in this study; however, species with less than 1000 ha of EO could also be categorized based on the criterion of an AO of less than 1,000 ha (criteria B2, for CR [48]) as well as species with an AO less than 50,000 ha (criteria B2, for EN [48]) because AO is always smaller than EO and is located within the EO polygon [48].
Population size was inferred for each species based on the estimates of area in m2 inhabited by one individual compiled by Bornschein et al. [24]. Based on estimates of the number of calling males [24], a sex ratio of one female per male [24] was assumed. In cases with distinct estimates of densities per species [24], the mean density was used. The mean area in which one individual per species can be found and its respective EO was then used to calculate the population size.
Data on EO, number of locations, population size, and threats of the species were integrated to evaluate and to categorize its conservation status according to the IUCN Red List and Criteria [48]. For the recognition of threats, data from the literature, personal field experience of the authors collected in the EO of 29 species, and information on land use, forest quality, and trends of deforestation over the previous 10 years were considered. For temporal trends in land use, the time series of satellite images of Google Earth Pro v. 7.1.4.1529 was analyzed.
In the treatment of the data in relation to the IUCN criteria, the flow chart presented in Figure 1 was used. Six pathways were developed beginning with the evaluation of the number of localities (one to two; three or more). If the species had up to two recorded localities, its altitudinal range was calculated. If an altitudinal range was not associated with the record, this prevented creating a polygon and estimating the EO. The species was then considered DD (pathway 3 of Figure 1). If the records were associated with altitudinal range, an EO was created based on the lower and upper altitudinal limits. It was not always possible to infer the EO without encompassing inland areas far west of the record and outside the assumed natural range, sometimes nearly reaching Argentina, which is clearly unrealistic. In these situations, the species were considered DD (pathway 2 of Figure 1). When there were up to two records associated with an altitudinal range that encompassed a realistic polygon for EO (as indicated), the status of the species was evaluated (pathway 1 of Figure 1). Further pathways related to the procedure can be observed in Figure 1.

3. Results

A total of 185 locality records representing all 36 currently recognized Brachycephalus species in addition to 32 Brachycephalus sp. were generated (Table 1). An unidentified Brachycephalus species represented one between two described species that could not be adequately identified (i.e., old museum material collected before certain species were described) as well as new species awaiting formal description. Hereafter, only the described species are analyzed, leaving any evaluations to their own descriptors. The EO for 26 species (Table 1) was estimated, comprising several highly restricted EOs as well as larger ones: 23.8 ha for B. fuscolineatus, 37.4 ha for B. coloratus, 38.8 ha for B. boticario, 41.4 ha for B. tridactylus, 56.8 ha for B. mirissimus (all from the B. pernix group), 143,325.0 ha for B. hermogenesi, 702,983.4 ha for B. didactylus, 3,021,786.1 ha for B. sulfuratus (B. didactylus group), and 1,792,535.1 ha for B. ephippium (B. ephippium group). The population sizes of eight species (Table 1) was also estimated. All were highly abundant with population sizes ranging from 78,344 individuals for B. mirissimus and 302,178,610 individuals for B. sulfuratus.
The main threat to the species of Brachycephalus is deforestation, affecting not only their EO but also other aspects of their biology, such as population size and individual health. Indeed, deforestation affects 20 species. Other species are under threat due to their small EO. Forests within EOs were converted into agricultural areas (e.g., for coffee and palm plantations - Archontophoenix alexandrae H. Wendl. & Drude), tree monocultures (Pinus spp. and Eucalyptus spp.), urban areas and, more frequently, pastures. Some species also have part of their EO flooded by dams (e.g., B. nodoterga) or affected by landslides (e.g., B. izecksohni). Fire, edge effects, timber harvest, grazing, and the invasion of exotic plants are impacts that reduce the quality of EO. For instance, deforestation and loss of habitat quality are important threats to B. mariaeterezae, whose type locality suffers from fire, grazing, and timber harvests. Fire and grazing substantially affect the quality of forests, even the cloud forests of B. quiririensis. The estimated population sizes were above those used in the IUCN criteria and therefore were not useful to rank the studied species regarding their conservation status.
The conservation status of all described species (Figure 2, Table 2) were determined. Twenty-one species (58.3% of all species) were classified as at risk of extinction: six as CR (28.6%), five as Endangered (EN; 23.8%), and 10 as Vulnerable (VU; 47.6%). Five species (13.9% of all species) were classified as non-threatened (= LC) and the remaining ten species (27.8%) as DD. The reduced EO (criteria B1) contributes to the ranking status of the conservation of 16 species associated with the number of locations (criteria B1a; 16 species), threats that reduce the area of EO (criteria B1b(i); 16 species), and quality of the area of EO (B1b(iii); 16 species; Table 2). B2 criteria were adopted for eight species (Table 2). For the B2 criteria, the number of locations (criteria B2a; eight species) and the threats that reduced the AO area (criteria B2b(ii); eight species) and quality (criteria B2b(iii); eight species) were also considered. Only one additional criterion (D2) for five species with less than 2000 ha of AO was used, and no threat could be assessed for this AO.

4. Discussion

Based on the assessments, the number of endangered species of Brachycephalus should increase from one to 21 (Table 2). This is a significant shift and poses the question regarding why only one species had been formally recognized as threatened until now (Table 2). There are two possible reasons: (1) a delay due to the fact that many species have only been recently described and (2) a resistance based on current policies of the MMA to change a species conservation status in a short period of time (see below). Twenty-two species have been described in the last 10 years, 14 of which are only known from their type locality, and there is a natural tendency to expect them to be more widely distributed; however, studies in recent years have been gradually revealing new species rather than new records of known species, and new records of already described species have not substantially altered their geographical distributions (e.g., [24,29,76,99]). For example, a new locality record for B. fuscolineatus published after its description increased its EO by just 0.19 ha [24], and this species still has the smallest estimated EO for any Brachycephalus species (Table 2). Two new records of B. curupira (Table 1, [29]) double its EO, which remains small (= 4751.4 ha; Table 2). A new record of B. albolineatus published after its description [76] and two new localities included in Table 1 substantially extend its EO, but as in the case of B. curupira, this remains small (= 2784.4 ha; Table 2). A new record for B. nodoterga [99] did not change the EO of the species because it is located within its EO polygon. Brachycephalus was not found in 13 localities from southern São Paulo to northeastern Santa Catarina with an altitude comparable to other localities where Brachycephalus populations were present. Overall, the reduced geographical distributions of Brachycephalus is the rule for the montane species of the genus, i.e., the B. ephippium and B. pernix groups [23]. Brachycephalus ephippium is the only exception of a montane species group with a large EO, but it is expected that some, if not all, populations may be identified as distinct species in future studies [23].
With respect to the resistance to incorporate drastic changes into the official number of endangered Brachycephalus species, this proposal is warranted despite the current policy of the MMA indicated. The MMA joined several international agreements that set targets for the conservation of the country’s threatened biodiversity, and these efforts have been implemented in the successful execution of National Action Plans for the Conservation of Brazilian Endangered Species (Planos de Ação Nacional - PANs). The national scientific community and the MMA have been working together to list threats and conservation actions to all threatened species of the country and to review and to monitor these actions annually. This is possibly the reason that the MMA tended to prefer moving forward with conservation strategies of species that are already listed as threatened rather than revising the list. The effort to prioritize conservation initiatives prior to substantial updates to the list of endangered species is recognized, but the need for MMA to revise the list and to recognize the species listed in this article as threatened is also acknowledged given that they are not yet legally protected.
The most prevalent threat to Brachycephalus is deforestation, much of which is no longer done with heavy machinery and chainsaws. Recently, deforestation in the Atlantic Rainforest has become more subtle and involves the selective removal of trees and shrubs, particularly through inconspicuous strategies, such as bark girdling, which leads to the opening of the canopy and an increased tendency for wind to knock down additional trees. These actions are deliberately conducted a few meters into the forest edge to avoid detection by environmental inspectors. This type of deforestation has been carried out at an alarming rate in Paraná and in the northeast of Santa Catarina for at least 25 years to cultivate bananas, and more recently, to cultivate palm trees (Archontophoenix alexandrae). Deforestation for agricultural activities could also result in soil contamination, affecting species that depend on specific microhabitats and that have permeable skin [121]. Finally, deforestation could exacerbate edge effects, altering microhabitats and microclimatic conditions, which changes sunlight exposure, soil moisture, and plant species composition in the edges [15]. Indeed, B. fuscolineatus was not encountered in forest edges but only in more nuclear vegetation [24].
Deforestation in lowlands can lead to a decrease in the altitudinal distribution of cloud forests [122], potentially shifting the distributions of montane species of Brachycephalus to higher altitudes. This possibility of altitudinal species displacement could also be driven by climate change [23]. In tropical forests, temperatures can vary from 0.4 °C to 0.7 °C per 100 m altitude variation [123]. The thermal variation in the altitudinal gradient in a site with the occurrence of Brachycephalus (B. pernix) was determined as 0.56 °C of the reduction every 100 m of altitude [124]. A difference in the precipitation levels at this site was also evaluated, with an increase of 40 mm in mean annual precipitation every 100 m at elevation [124]. Studies on litter anurans of the Atlantic Rainforest, including Brachycephalus, have shown that population densities are particularly affected by air humidity, air temperature, and altitude [52,55,78,90]. This climatic dependence and its relationship with the altitude gradient raises concerns for the long-term conservation of Brachycephalus species that occur in mountains with a restricted altitudinal amplitude.
Lowering the category of threat for B. pernix from CR [45] to VU (Table 2) is proposed. The effects of trampling and timber harvests by tourists in the type locality of the species are likely to be minor, which is entirely distributed within a protected area (Parque Estadual Serra da Baitaca; Table S1). There is a threat of fire in part of the EO of the species, but the vegetation cover is regenerating well in this area after many years of management by volunteer mountaineers, reducing fire susceptibility.
It is recognized that there is some level of subjectivity to apply EO and the number of locations of threatened species. This is because each parameter shows some overlap between EN and VU categories. Brachycephalus hermogenesi and B. nodoterga fit the EN category, but both are recommended to be considered for the VU category because part of their EO is in protected areas (Table S1).
The presence of threatened Brachycephalus in protected areas is a useful tool to rank the species for which conservation actions are more urgent. In Table S1, 10 species without records in protected areas are recognized with three classified as CR (B. boticario, B. mirissimus, and B. quiririensis), two as EN (B. actaeus and B. leopardus), one as VU (B. albolineatus), and four as DD (B. atelopoide, B. auroguttatus, B. bufonoides, B. leopardus, and B. verrucosus). There are no known living populations of two species (B. atelopoide and B. bufonoides). The remaining eight species belonging to the B. pernix group occur in southern Paraná (B. leopardus) and Santa Catarina (B. actaeus, B. albolineatus, B. auroguttatus, B. boticario, B. mirissimus, B. quiririensis, and B. verrucosus). Also, it is argued that DD species need special attention to direct further studies to complete adequate assessments of their conservation status as soon as possible.
Santa Catarina stands out as the state in which emergency conservation actions should converge. Creating protected areas is an important way to protect species, however, the conservation of the top three priority species would require the creation of three new protected areas. A protected area for the CR B. quiririensis could already house other species of Brachycephalus that are not in any reserve, namely B. leopardus (EN) and B. auroguttatus (DD). Nonetheless, to be effective, a protected area would first require the expropriation of the land in addition to management actions aimed at recovering forest quality. Given that there are dozens of protected areas waiting for expropriation, this path to conservation does not seem likely at the moment. Private protected areas are an alternative (e.g., Reserva Particular do Patrimônio Natural—RPPN), and some of them already protect two species of Brachycephalus (B. mariaeterezae and B. tridactylus; Table 1 and Table S1). This is the most stable category of protected areas in Brazil and cannot be undone; however, one aspect that does not stimulate the creation of more private protected areas is the lack of government incentives to private owners, except for exemption from territorial taxes. There is an impediment to transferring public financial resources to private persons, even if they are addressing conservation measures.
The conservation of Brachycephalus should also include alternatives to the creation of protected areas. One approach would be to lease land with the occurrence of threatened Brachycephalus at a percentage of the regional value of production per hectare of mountainous lands, which would be an incentive for landowners to leave their land intact. This must be governed by a renewable contract. For this strategy to be put into practice, it is vital to attract international resources. It would also be interesting to attract additional resources of the lease value to promote environmental recovery. The management of invasive alien species, both plants and animals, is unfortunately incipient in Brazil due to the high involved costs, thus discouraging the proposition of new management projects. The state of Santa Catarina has emphasized its concern with the conservation of microendemic anurans in its region, creating a specific program for this purpose (Portaria Instituto Estadual do Meio Ambiente - IMA N° 283/2018 - 19/12/2018). This is an interesting effort that can put actions discussed into practice and can also result in other effective and innovative actions for the conservation of Brachycephalus in Santa Catarina. In the long term, successful practices can be replicated in other regions of Brazil.

5. Conclusions

Advances in knowledge regarding the geographical distribution of the Brachycephalus species have confirmed that they are in fact restricted, and this restriction is the reason for classifying 58% of the species of the genus as threatened according to IUCN criteria. Restricted geographical distributions should be considered an attribute of the species of the Brachycephalus montane groups. This coincides with the tendency of species with small ranges to be geographically concentrated and disproportionately under the threat of extinction [125] as well as with the tendency of newly described species to be more threatened than those described earlier [3]. With an increased understanding of the nature of most Brachycephalus species as microendemic species, international (IUCN) and national (MMA) agencies might be more likely to update their conservation status based on this proposal. Furthermore, Brazil has the highest amphibian richness in the world and the highest description of new species in recent years, but it is one of the countries with the lowest update rates of conservation status [3].
Deforestation and loss of habitat quality impact almost all species of Brachycephalus (22 species). Species of the genus are locally highly abundant, but they respond in density and geographical distribution to temperature and humidity [23,24,27], which vary along the altitudinal gradient [122,124]. Climate change can influence climatic conditions along the altitudinal gradient, confining the distribution of species even further to higher altitudes in the future.
The common action to protect endangered species in Brazil is to create protected areas. The creation of a new protected area in southern Paraná (Serra do Araçatuba) and adjacent to Santa Catarina (Serra do Quiriri) is proposed, but only because it would protect three species (B. quiririensis—CR, B. leopardus—EN, and B. auroguttatus—DD). In the marshes and grasslands associated with the forest of occurrence of these three Brachycephalus species is another endangered frog, Melanophryniscus biancae, which is a candidate for EN [126,127]. One reserve including the distribution of these four species would have about 11,000 ha—6,000 ha of forests, and 5,000 ha of grasslands [126]—and would also protect the springs of important rivers, such as the Negro, Cubatão, and Pirabeiraba. The creation of several other protected areas to safeguard the remaining threatened species without occurrence in reserves is impractical in the current Brazilian economic scenario. A program to lease strategic private land for owners to keep them intact with the support of international resources is a possibility for the conservation of the species in the short and medium term.

Supplementary Materials

The following are available online at https://www.mdpi.com/1424-2818/11/9/150/s1, Table S1: Records and distribution of the extent of occurrence (EO) of the species of Brachycephalus in relation to protected areas (PA).

Author Contributions

Conceptualization, M.R.B.; methodology, M.R.B.; formal analysis, M.R.B.; investigation, M.R.B.; data curation, M.R.B.; writing—original draft preparation, M.R.B. and L.T.; writing—review and editing, M.R.B., L.T. and M.R.P.

Funding

Field researches was partially funded by Fundação Grupo Boticário de Proteção à Natureza (through grants 0895_20111 and A0010_2014, conducted by Mater Natura—Instituto de Estudos Ambientais) and National Geographic Society (through the grant EC–50722R-18 to L.T.) L.T. was supported through a grant from CAPES / Reitoria and M.R.P. through a grant from CNPq/MCT (571334/2008–3).

Acknowledgments

Luiz F. Ribeiro provided valuable assistance during the field work. We thank two anonymous reviewers for comments on the manuscript.

Conflicts of Interest

The authors declare no conflict of interest. The funders had no role in the design of the study, the collection, analyses, or interpretation of data, the writing of the manuscript, or the decision to publish the results.

References

  1. Frost, D.R. Amphibian Species of the World: An Online Reference, Version 6.0. 2019. Available online: http://research.amnh.org/herpetology/amphibia/index.html (accessed on 22 June 2019).
  2. Köhler, K.; Vieites, D.R.; Bonett, R.M.; Garcia, F.H.; Glaw, F.; Steinke, D.; Vences, M. New Amphibians and Global Conservations: A boost in species discoveries in a highly endangered vertebrate group. BioScience 2005, 55, 693–696. [Google Scholar] [CrossRef]
  3. Tapley, B.; Michaels, C.J.; Gumbs, R.; Böhm, M.; Luedtke, J.; Pearce-Kelly, P.; Rowley, J.J.L. The disparity between species description and conservation assessment: A case study in taxa with high rates of species discovery. Biol. Conserv. 2018, 220, 209–214. [Google Scholar] [CrossRef]
  4. Wake, D.B.; Vredenburg, V.T. Are we in the midst of the sixth mass extinction? A view from the world of amphibians. Proc. Natl. Acad. Sci. USA 2008, 105, 11466–11473. [Google Scholar] [CrossRef] [PubMed][Green Version]
  5. IUCN. The IUCN Red List of Threatened Species. Available online: https://www.iucnredlist.org (accessed on 28 June 2019).
  6. Stuart, S.; Chanson, J.; Cox, N. Status and trends of amphibian declines and extinctions worldwide. Science 2004, 306, 1783–1786. [Google Scholar] [CrossRef] [PubMed]
  7. Skerrat, L.F.; Berger, L.; Speare, R.; Cashins, S.; McDonald, K.R. Spread of Chytridiomycosis Has Caused the Rapid Global Decline and Extinction of Frogs. EcoHealth 2007, 4, 126. [Google Scholar] [CrossRef]
  8. Stuart, S.N.; Hoffmann, M.; Chanson, J.S.; Cox, N.A.; Berridge, R.J.; Ramani, P.; Young, B.E. Threatened Amphibians of the World; Lynx Editions: Barcelona, Spain, 2008; pp. 1–151. [Google Scholar]
  9. Lips, K.P.R.; Burrowes, P.A.; Mendelson, J.R., III; Parra-Olea, G. Amphibian declines in Latins America: A synthesis. Biotropica 2005, 37, 222–226. [Google Scholar] [CrossRef]
  10. Ruland, F.; Jeschke, J.M. Threat-dependent traits of endangered frogs. Biol. Conserv. 2017, 206, 310–313. [Google Scholar] [CrossRef]
  11. Ruggeri, J.; Ribeiro, L.P.; Pontes, M.R.; Toffolo, C.; Candido, M.; Carriero, M.M.; Zanella, N.; Sousa, R.L.M.; Toledo, L.F. First case of wild amphibians infected with Ranavirus in Brazil. J. Wildlife Dis. 2019, 55. Preprint. [Google Scholar]
  12. Myers, N.; Mittermeier, R.A.; Mittermeier, C.G.; Fonseca, G.A.B.; Kent, J. Biodiversity hotspots for conservation priorities. Nature 2000, 403, 853–858. [Google Scholar] [CrossRef] [PubMed]
  13. Dov-Por, F. Sooretema, the Atlantic Rain Forest of Brazil; SPB Academic Publishing: The Hague, The Netherlands, 1992; Volume 128. [Google Scholar]
  14. MMA. Mapa de Vegetação Nativa na Área de Aplicação da Lei no. 11.428/2006—Lei da Mata Atlântica (ano base 2009); Ministério do Meio Ambiente: Brasília, Brazil, 2015; pp. 1–85.
  15. Rezende, C.L.; Scarano, F.R.; Assad, E.D.; Joly, C.A.; Metzger, J.P.; Strassburg, B.B.N.; Tabarelli, M.; Fonseca, G.A.; Mittermeier, R.A. From hotspots to hopespot: An opportunity for the Brazilian Atlantic Forest. Perspect. Ecol. Conserv. 2018, 16, 204–214. [Google Scholar] [CrossRef]
  16. Ministério do Meio Ambiente. Livro Vermelho da Fauna Brasileira Ameaçada de Extinção—Sumário Executivo; Instituto Chico Mendes de Conservação da Biodiversidade—ICMBio: Brasília, Brazil, 2016; pp. 1–76.
  17. Izecksohn, E. Novo gênero e nova espécie de Brachycephalidae do Estado do Rio de Janeiro, Brasil. Boletim do Museu Nacional Zoologia 1971, 280, 1–12. [Google Scholar]
  18. Giaretta, A.A.; Sawaya, R.J. Second species of Psyllophryne (Anura: Brachycephalidae). Copeia 1998, 1998, 985–987. [Google Scholar] [CrossRef]
  19. Pombal, J.P., Jr. Oviposição e desenvolvimento de Brachycephalus ephippium (Spix) (Anura, Brachycephalidae). Rev. Brasil. Zool. 1999, 16, 967–976. [Google Scholar] [CrossRef]
  20. Napoli, M.F.; Caramaschi, U.; Cruz, C.A.G.; Dias, I.R. A new species of flea-toad, genus Brachycephalus Fitzinger (Amphibia: Anura: Brachycephalidae), from the Atlantic Rainforest of southern Bahia, Brazil. Zootaxa 2011, 2739, 33–40. [Google Scholar] [CrossRef]
  21. Yeh, J. The effect of miniaturized body size on skeletal morphology in frogs. Evolution 2002, 56, 628–641. [Google Scholar] [CrossRef] [PubMed]
  22. Schwartz, C.A.; de Souza Castro, M.; Pires Júnior, O.R.; Maciel, N.M.; Schwartz, E.N.F.; Sebben, A. Princípios bioativos da pele de anfíbios: Panorama atual e perspectivas. In Herpetologia no Brasil II; Nascimento, L.B., Oliveira, M.E., Eds.; Sociedade Brasileira de Herpetologia: Belo Horizonte, Brasil, 2007; pp. 146–168. [Google Scholar]
  23. Bornschein, M.R.; Firkowski, C.R.; Belmonte-Lopes, R.; Corrêa, L.; Ribeiro, L.F.; Morato, S.A.A.; Antoniazzi, R.L., Jr.; Reinert, B.L.; Meyer, A.L.S.; Cini, F.A.; et al. Geographical and altitudinal distribution of Brachycephalus (Anura: Brachycephalidae) endemic to the Brazilian Atlantic Rainforest. PeerJ 2016, 4, e2490. [Google Scholar] [CrossRef] [PubMed]
  24. Bornschein, M.R.; Teixeira, L.; Ribeiro, L.F. New record of Brachycephalus fuscolineatus Pie, Bornschein, Firkowski, Belmonte-Lopes & Ribeiro, 2015 (Anura, Brachycephalidae) from Santa Catarina state, Brazil. Check List 2019, 15, 379–385. [Google Scholar]
  25. Pie, M.R.; Ribeiro, L.F.; Confetti, A.E.; Nadaline, M.J.; Bornschein, M.R. A new species of Brachycephalus (Anura: Brachycephalidae) from southern Brazil. PeerJ 2018, 6, e5683. [Google Scholar] [CrossRef] [PubMed]
  26. Ribeiro, L.F.; Bornschein, M.R.; Belmonte-Lopes, R.; Firkowski, C.R.; Morato, S.A.A.; Pie, M.R. Seven new microendemic species of Brachycephalus (Anura: Brachycephalidae) from southern Brazil. PeerJ 2015, 3, e1011. [Google Scholar] [CrossRef] [PubMed]
  27. Pie, M.R.; Meyer, A.L.S.; Firkowski, C.R.; Ribeiro, L.F.; Bornschein, M.R. Understanding the mechanisms underlying the distribution of microendemic montane frogs (Brachycephalus spp., Terrarana: Brachycephalidae) in the Brazilian Atlantic Rainforest. Ecol. Model. 2013, 250, 165–176. [Google Scholar] [CrossRef]
  28. Firkowski, C.R.; Bornschein, M.R.; Ribeiro, L.F.; Pie, M.R. Species delimitation, phylogeny and evolutionary demography of co-distributed, montane frogs in the southern Brazilian Atlantic Forest. Mol. Phylogenet. Evol. 2016, 100, 345–360. [Google Scholar] [CrossRef] [PubMed]
  29. Pie, M.R.; Faircloth, B.C.; Bornschein, M.R.; McComarck, J.E. Phylogenomics of montane frogs of the Brazilian Atlantic Forest is consistent with isolation in sky islands followed by climatic stability. Biol. J. Linn. Soc. 2018, 125, 72–82. [Google Scholar] [CrossRef]
  30. Pombal, J.P., Jr. A posição taxonômica das “variedades” de Brachycephalus ephippium (Spix, 1824) descritas por Miranda-Ribeiro, 1920 (Amphibia, Anura, Brachycephalidae). Bol. Mus. Nac. Zool. 2010, 526, 1–12. [Google Scholar]
  31. Silvano, D.; Heyer, R.; Caramaschi, U. Brachycephalus nodoterga. The IUCN Red List of Threatened Species 2004: e.T54454A11149387. Available online: https://www.iucnredlist.org/species/54454/11149387 (accessed on 13 July 2019).
  32. Silvano, D.; Garcia, P.; Segalla, M.V. Brachycephalus pernix. The IUCN Red List of Threatened Species 2004, e.T54455A11149530. Available online: https://www.iucnredlist.org/species/54455/11149530 (accessed on 13 July 2019).
  33. Silvano, D.; Caramaschi, U. Brachycephalus hermogenesi. The IUCN Red List of Threatened Species 2010, e.T29487A9501270. Available online: https://www.iucnredlist.org/species/29487/9501270 (accessed on 13 July 2019).
  34. Angulo, A. Brachycephalus ferruginus. The IUCN Red List of Threatened Species 2006, e.T135912A4229152. Available online: https://www.iucnredlist.org/species/135912/4220152 (accessed on 13 July 2019).
  35. Angulo, A. Brachycephalus alipioi. The IUCN Red List of Threatened Species 2008, e.T135774A4199662. Available online: https://www.iucnredlist.org/species/135774/4199662 (accessed on 13 July 2019).
  36. Angulo, A. Brachycephalus pombali. The IUCN Red List of Threatened Species 2008, e.T135830A4208137. Available online: https://www.iucnredlist.org/species/135830/4208137 (accessed on 13 July 2019).
  37. Caramaschi, U.; Carvalho-e-Silva, S.P. Brachycephalus vertebralis. The IUCN Red List of Threatened Species 2004, e.T54456A11134718. Available online: https://www.iucnredlist.org/species/54456/11134718 (accessed on 13 July 2019).
  38. Sluys, M.V.; Rocha, C.F. Brachycephalus ephippium. The IUCN Red List of Threatened Species 2010: e.T54453A11149233. Available online: https://www.iucnredlist.org/species/54453/11149233 (accessed on 13 July 2019).
  39. Stuart, S. Brachycephalus brunneus. The IUCN Red List of Threatened Species 2006, e.T61745A12553521. Available online: https://www.iucnredlist.org/species/61746/12553521 (accessed on 13 July 2019).
  40. Stuart, S. Brachycephalus izecksohni. The IUCN Red List of Threatened Species 2006, e.T61747A12553638. Available online: https://www.iucnredlist.org/species/61747/12553638 (accessed on 13 July 2019).
  41. Telles, A.M.; Carvalho-e-Silva, S.P. Brachycephalus didactylus. The IUCN Red List of Threatened Species 2004, e.T54452A11148997. Available online: https://www.iucnredlist.org/species/54452/11148997 (accessed on 13 July 2019).
  42. Haddad, C.F.B.; Machado, I.F.; Giovanelli, J.G.R.; Bataus, Y.S.L.; Ublig, V.M.; Batista, F.R.Q.; Cruz, C.A.G.; Conte, C.E.; Zank, C.; Strüsmann, C.; et al. Avaliação do Risco de Extinção de Brachycephalus nodoterga Miranda-Ribeiro, 1920. In Processo de Avaliação do Risco de Extinção da Fauna Brasileira; Instituto Chico Mentes de conservação da Biodiversidade—ICMbio: Brasília, Brazil, 2016. [Google Scholar]
  43. Haddad, C.F.B.; Machado, I.F.; Giovanelli, J.G.R.; Bataus, Y.S.L.; Uhlig, V.M.; Batista, F.R.Q.; Cruz, C.A.G.; Conte, C.E.; Zank, C.; Strüsmann, C.; et al. Avaliação do Risco de Extinção de Brachycephalus alipioi Pombal & Gasparini, 2006. In Processo de Avaliação do Risco de Extinção da Fauna Brasileira; Instituto Chico Mentes de conservação da Biodiversidade—ICMbio: Brasília, Brazil, 2016. [Google Scholar]
  44. Haddad, C.F.B.; Machado, I.F.; Giovanelli, J.G.R.; Bataus, Y.S.L.; Uhlig, V.M.; Batista, F.R.Q.; Cruz, C.A.G.; Conte, C.E.; Zank, C.; Strüsmann, C.; et al. Avaliação do Risco de Extinção de Brachycephalus vertebralis Pombal, 2001. In Processo de Avaliação do Risco de Extinção da Fauna Brasileira; Instituto Chico Mentes de conservação da Biodiversidade—ICMbio: Brasília, Brazil, 2016. [Google Scholar]
  45. Haddad, C.F.B.; Segalla, M.V.; Bataus, Y.S.L.; Uhlig, V.M.; Batista, F.R.Q.; Garda, A.; Hudson, A.A.; Cruz, C.A.G.; Strüsmann, C.; Brasileiro, C.A.; et al. Avaliação do Risco de Extinção de Brachycephalus pernix Pombal, Wistuba & Bornschein, 1998. In Processo de Avaliação do Risco de Extinção da Fauna Brasileira; Instituto Chico Mentes de conservação da Biodiversidade—ICMbio: Brasília, Brazil, 2016. [Google Scholar]
  46. Bland, L.M.; Colle, B.; Orme, C.D.L.; Bielby, J. Data uncertainty and the selectivity of extinction risk in freshwater invertebrates. Divers. Distrib. 2012, 18, 1211–1220. [Google Scholar] [CrossRef]
  47. Morais, A.R.; Siqueira, M.N.; Lemes, P.; Maciel, N.M.; De Marco, P.; Brito, D. Unraveling the conservations status of Data Deficient species. Biol. Conserv. 2013, 166, 98–102. [Google Scholar] [CrossRef]
  48. IUCN. IUCN Red List Categories and Criteria: Version 3.1., 2nd ed.; International Union for Conservation of Nature—IUCN: Gland, Switzerland; Cambridge, UK, 2012. [Google Scholar]
  49. Reinert, B.L.; Bornschein, M.R.; Firkowski, C. Distribuição, tamanho populacional, hábitat e conservação do bicudinho-do-brejo Stymphalornis acutirostris Bornschein, Reinert e Teixeira, 1995 (Thamnophilidae). Rev. Bras. Ornitol. 2007, 15, 493–519. [Google Scholar]
  50. Ribeiro, L.F.; Blackburn, D.C.; Stanley, E.L.; Pie, M.R.; Bornschein, M.R. Two new species of the Brachycephalus pernix group (Anura: Brachycephalidae) from the state of Paraná, southern Brazil. PeerJ 2017, 5, e3603. [Google Scholar] [CrossRef] [PubMed]
  51. De Oliveira, J.C.F.; Coco, L.; Pagotto, R.V.; Pralon, E.; Vrcibradic, D.; Pombal, J.P., Jr.; Rocha, C.F.D. Amphibia, Anura, Brachycephalus didactylus (Izecksohn, 1971) and Zachaenus parvulus (Girard, 1853): Distribution extension. Check List 2012, 8, 242–244. [Google Scholar] [CrossRef]
  52. Oliveira, J.C.F.; Pralon, E.; Coco, L.; Pagotto, R.V.; Rocha, C.F.D. Environmental humidity and leaf-litter depth affecting ecological parameters of a leaf-litter frog community in an Atlantic Rainforest area. J. Nat. Hist. 2013, 47, 2115–2124. [Google Scholar] [CrossRef]
  53. Siqueira, C.C.; Vrcibradic, D.; Almeida-Gomes, M.; Borges, V.N.T., Jr.; Almeida-Santos, P.; Almeida-Santos, M.; Ariani, C.V.; Guedes, D.M.; Goyannes-Araújo, P.; Dorigo, T.A.; et al. Density and richness of the leaf litter frogs of an Atlantic Rainforest area in Serra dos Órgãos, Rio de Janeiro State, Brazil. Zoologia 2009, 26, 97–102. [Google Scholar] [CrossRef]
  54. Almeida-Santos, M.; Siqueira, C.C.; Van Sluys, M.; Rocha, C.D.F. Ecology of the Brazilian flea frog Brachycephalus didactylus (Terrana: Brachycephalidae). J. Hepertol. 2011, 45, 251–255. [Google Scholar]
  55. Siqueira, C.C.; Vrcibradic, D.; Nogueira-Costa, P.; Martins, A.R.; Dantas, L.; Gomes, V.L.R.; Bergallo, H.G.; Rocha, C.F.D. Environmental parameters affecting the structure of leaf-litter frog (Amphibia: Anura) communities in tropical forests: A case study from an Atlantic Rainforest area in southeastern Brazil. Zoologia 2014, 31, 147–152. [Google Scholar] [CrossRef]
  56. Carvalho-e-Silva, A.M.T.; Silva, G.R.; Carvalho-e-Silva, S.P. Anuros da Reserva Rio das Pedras, Mangaratiba, RJ, Brasil. Biota Neotrop. 2008, 8, 199–209. [Google Scholar] [CrossRef][Green Version]
  57. Rocha, C.F.D.; Vrcibradic, D.; Kiefer, M.C.; Almeida-Gomes, M.; Borges-Junior, V.N.T.; Menezes, V.A.; Ariani, C.V.; Pontes, J.A.L.; Goyannes-Araújo, P.; Marra, R.V.; et al. The leaf-litter frog community from Reserva Rio das Pedras, Mangaratiba, Rio de Janeiro State, Southeastern Brazil: Species richness, composition and densities. North West. J. Zool. 2013, 9, 151–156. [Google Scholar]
  58. Pombal, J.P., Jr. A new species of Brachycephalus (Anura: Brachycephalidae) from Atlantic Forest of southeastern Brazil. Amphib. Reptil. 2001, 22, 179–185. [Google Scholar] [CrossRef]
  59. Ribeiro, L.F.; Alves, A.C.R.; Haddad, C.F.B. Two new species of Brachycephalus Günther, 1858 from the state of Paraná, southern Brazil (Amphibia, Anura, Brachycephalidae). Bol. Mus. Nac. Zool. 2005, 519, 1–18. [Google Scholar]
  60. Alves, A.C.R.; Ribeiro, L.F.; Haddad, C.F.B.; Reis, S.F. Two new species of Brachycephalus (Anura: Brachycephalidae) from the Atlantic Forest in Paraná State, southern Brasil. Hepertologica 2006, 62, 221–233. [Google Scholar] [CrossRef]
  61. Alves, A.C.R.; Sawaya, R.J.; Reis, S.F.; Haddad, C.F.B. New species of Brachycephalus (Anura: Brachycephalidae) from the Atlantic Rain Forest in São Paulo State, Southeastern Brazil. J. Hepertol. 2009, 43, 212–219. [Google Scholar] [CrossRef]
  62. Da Silva, H.R.; Campos, L.A.; Sebben, A. The auditory region of Brachycephalus and its bearing on the monophyly of the genus (Anura: Brachycephalidae). Zootaxa 2007, 1422, 59–68. [Google Scholar] [CrossRef]
  63. Verdade, V.K.; Rodrigues, M.T.; Cassimiro, J.; Pavan, D.; Liou, N.; Lange, M. Advertisement call, vocal activity, and geographic distribution of Brachycephalus hermogenesi (Giaretta and Sawaya, 1998) (Anura, Brachycephalidae). J. Herpetol. 2008, 42, 542–549. [Google Scholar] [CrossRef]
  64. Clemente-Carvalho, R.G.B.; Antoniazzi, M.M.; Jared, C.; Haddad, C.F.B.; Alvez, A.C.R.; Rocha, H.S.; Pereira, G.R.; Oliveira, D.F.; Lopes, R.T.; Reis, S.F. Hyperossification in miniaturized toadlets of the genus Brachycephalus (Amphibia: Anura: Brachycephalidae): Microscopic structure and macroscopic patterns of variation. J. Morphol. 2009, 270, 1285–1295. [Google Scholar] [CrossRef] [PubMed]
  65. Campos, L.A. Sistemática Filogenética do Gênero Brachycephalus Ftzinger, 1826 (Anura Brachycephalidae) Com Base Em Dados Morfológicos. Ph.D. Thesis, Universidade de Brasília, Brasília, Brazil, 2011. [Google Scholar]
  66. Pombal, J.P., Jr.; Izecksohn, E. Uma nova espécie de Brachycephalus (Anura, Brachycephalidae) do Estado do Rio de Janeiro. Pap. Av. Zool. 2011, 51, 443–451. [Google Scholar] [CrossRef]
  67. Siqueira, C.C.; Vrcibradic, D.; Dorigo, T.A.; Rocha, C.F.D. Anurans from two high-elevation areas of Atlantic Forest in the state of Rio de Janeiro, Brazil. Zoologia 2011, 28, 457–464. [Google Scholar] [CrossRef]
  68. Rocha, C.F.D.; Van Sluys, M.; Alves, M.A.S.; Bergallo, H.G.; Vrcibradic, D. Activity of leaf-litter frogs: When should frogs be sampled? J. Herpetol. 2000, 34, 285–287. [Google Scholar] [CrossRef]
  69. Rocha, C.F.D.; Van Sluys, M.; Alves, M.A.S.; Bergallo, H.G.; Vrcibradic, D. Estimates of forest floor litter frog communities: A comparison of two methods. Austral. Ecol. 2001, 26, 14–21. [Google Scholar] [CrossRef]
  70. Van Sluys, M.; Vrcibradic, D.; Alves, M.A.S.; Bergallo, H.G.; Rocha, C.F.D. Ecological parameters of the leaf-litter frog community of an Atlantic Rainforest area at Ilha Grande, Rio de Janeiro state, Brazil. Austral. Ecol. 2007, 32, 254–260. [Google Scholar] [CrossRef]
  71. Pimenta, B.V.S.; Bérnils, R.S.; Pombal, J.P., Jr. Amphibia, Anura, Brachycephalidae, Brachycephalus hermogenesi: Filling gap and geographic distribution map. Check List 2007, 3, 277–279. [Google Scholar] [CrossRef]
  72. Condez, T.H.; Monteiro, J.P.C.; Comitti, E.J.; Garcia, P.C.A.; Amaral, I.B.; Haddad, C.F.B. A new species of flea-toad (Anura: Brachycephalidae) from southern Atlantic Forest, Brazil. Zootaxa 2016, 4083, 40–56. [Google Scholar] [CrossRef]
  73. Firkowski, C.R. Diversification and microendemism in montane refugia from the Brazilian Atlantic Forest. Master’s Thesis, Universidade Federal do Paraná, Curitiba, Brazil, 2013. [Google Scholar]
  74. Bornschein, M.R.; Ribeiro, L.F.; Blackburn, D.C.; Stanley, E.L.; Pie, M.R. A new species of Brachycephalus (Anura: Brachycephalidae) from Santa Catarina, southern Brazil. PeerJ 2016, 4, e2629. [Google Scholar] [CrossRef]
  75. Monteiro, J.P.C.; Condez, T.H.; Garcia, P.C.A.; Comitti, E.J.; Amaral, I.B.; Haddad, C.F.B. A new species of Brachycephalus (Anura, Brachycephalidae) from the coast of Santa Catarina State, southern Atlantic Forest, Brazil. Zootaxa 2018, 4407, 483–505. [Google Scholar] [CrossRef]
  76. Teixeira, L.; Ribeiro, L.F.; Côrrea, L.; Confetti, A.E.; Pie, M.R.; Bornschein, M.R. A second record of the recently described Brachycephalus albolineatus Bornschein, Ribeiro, Blackburn, Stanley & Pie, 2016 (Anura, Brachycephalidae). Check List 2018, 14, 1013–1016. [Google Scholar]
  77. Pereira, M.S.; Candaten, A.; Milani, D.; Oliveira, F.B.; Gardelin, J.; Rocha, C.F.D.; Vrcibradic, D. Geographic distribution: Brachycephalus hermogenesi. Herpetol. Rev. 2010, 41, 506. [Google Scholar]
  78. Santos-Pereira, M.; Candaten, A.; Milani, D.; Oliveira, F.B.; Gardelin, J.; Rocha, C.F.D. Seasonal variation in the leaf-litter frog community (Amphibia: Anura) from an Atlantic Forest area in the Salto Morato Natural Reserve, southern Brazil. Zoologia 2011, 28, 755–761. [Google Scholar] [CrossRef]
  79. Santos-Pereira, M.; Milani, D.; Barata-Bittencourt, L.F.; Iapp, T.M.; Rocha, C.F.D. Anuran species of the Salto Morato Nature Reserve in Paraná, southern Brazil: Review of the species list. Check List 2016, 12, 1907. [Google Scholar] [CrossRef]
  80. Pombal, J.P., Jr.; Gasparini, J.L. A new Brachycephalus (Anura: Brachycephalidae) from the Atlantic Rainforest of Espírito Santo, southeastern Brazil. S. Am. J. Herpet. 2006, 1, 87–93. [Google Scholar] [CrossRef]
  81. Clemente-Carvalho, R.G.B.; Klaczko, J.; Perez, S.R.; Alves, A.C.R.; Haddad, C.F.B.; Reis, S.F. Molecular phylogenetic relationships and phenotypic diversity in miniaturized toadlets, genus Brachycephalus (Amphibia: Anura: Brahycephalidae). Mol. Phylogenet. Evol. 2011, 61, 79–89. [Google Scholar] [CrossRef]
  82. Clemente-Carvalho, R.B.G.; Giaretta, A.A.; Condez, T.H.; Haddad, C.F.B.; Reis, S.F. A new species of miniaturized toadlet, genus Brachycephalus (Anura: Brachycephalidae), from the Atlantic Forest of southeastern Brazil. Herpetologica 2012, 68, 365–374. [Google Scholar] [CrossRef]
  83. Miranda-Ribeiro, A. Os Brachycephalideos do Museu Paulista (com tres especies novas). Rev. Mus. Paulista 1920, 12, 306–318. [Google Scholar]
  84. Condez, T.H.; Clemente-Carvalho, R.B.G.; Haddad, C.F.B.; Reis, S.F. A new species of Brachycephalus (Anura: Brachycephalidae) from the highlands of the Atlantic Forest, southeastern Brazil. Herpetologica 2014, 70, 89–99. [Google Scholar] [CrossRef]
  85. Moura, M.R.; Motta, A.P.; Fernandes, V.D.; Feio, R.N. Herpetofauna from Serra do Brigadeiro, an Atlantic Forest remain in the state of Minas Gerais, southeastern Brazil. Biota Neotrop. 2012, 12, 209–235. [Google Scholar] [CrossRef]
  86. Guimarães, C.S.; Luz, S.; Rocha, P.C.; Feio, R.N. The dark side of pumpkin toadlet: A new species of Brachycephalus (Anura: Brachycephalidae) from Serra do Brigadeiro, southeastern Brazil. Zootaxa 2017, 4258, 327–344. [Google Scholar] [CrossRef]
  87. Pombal, J.P., Jr.; Sazima, I.; Haddad, C.F.B. Breeding behavior of the pumpkin toadlet, Brachycephalus ephippium (Brachycephalidae). J. Herpetol. 1994, 28, 516–519. [Google Scholar] [CrossRef]
  88. Giaretta, A.A. Diversidade e Densidade de Anuros de Serapilheira Num Gradiente Altitudinal na Mata Atlântica Costeira. Ph.D. Thesis, Universidade Estadual de Campinas, Campinas, Brazil, 1999. [Google Scholar]
  89. Giaretta, A.A.; Facure, K.G.; Sawaya, R.J.; Meyer, J.H.M.; Chemin, N. Diversity and abundance of litter frogs in a montane forest of Southeastern Brazil: Seasonal and altitudinal changes. Biotropica 1999, 31, 669–674. [Google Scholar] [CrossRef]
  90. Giaretta, A.A.; Sawaya, R.J.; Machado, G.; Araújo, M.S.; Facure, K.G.; Medeiros, H.F.; Nunes, R. Diversity and abundance of litter frogs at altitudinal sites at Serra do Japi, Southeastern Brazil. Rev. Brasil. Zool. 1997, 14, 341–346. [Google Scholar] [CrossRef]
  91. Clemente-Carvalho, R.B.G.; Monteiro, L.R.; Bonato, V.; Rocha, H.S.; Pereira, G.R.; Oliveira, D.F.; Lopes, R.T.; Haddad, C.F.B.; Martins, E.G.; Reis, S.F. Geographic variation in cranial shape in the Pumpkin Toadlet (Brachycephalus ephippium): A geometric analysis. J. Herpetol. 2008, 42, 176–185. [Google Scholar] [CrossRef]
  92. Clemente-Carvalho, R.G.B.; Alves, A.C.R.; Perez, S.I.; Haddad, C.F.B.; Reis, S.F. Morphological and molecular variation in the Pumpkin Toadlet Branchycephalus ephippium (Anura: Brachycephalidae). J. Herpetol. 2011, 45, 94–99. [Google Scholar] [CrossRef]
  93. Zaher, H.; Aguiar, E.; Pombal, J.P. Paratelmatobius gaigeae (Cochran, 1938) rediscovered (Amphibia, Anura, Leptodactylidae). Arquiv. Mus. Nac. 2005, 63, 321–328. [Google Scholar]
  94. Dixo, M.; Verdade, V.K. Herpetofauna de serrapilheira da Reserva Florestal de Morro Grande, Cotia (SP). Biota Neotrop. 2006, 6, 1–20. [Google Scholar] [CrossRef]
  95. Siqueira, C.C.; Vrcibradic, D.; Rocha, C.F.D. Altitudinal records of data-deficient and threatened frog species from the Atlantic Rainforest of the Serra dos Órgãos mountains, in southeastern Brazil. Braz. J. Biol. 2013, 73, 229–230. [Google Scholar] [CrossRef]
  96. Dorigo, T.A.; Siqueira, C.C.; Vrcibradic, D.; Maia-Carneiro, T.; Almeida-Santos, M.; Rocha, C.F.D. Ecological aspects of the pumpkin toadlet, Brachycephalus garbeanus Miranda-Ribeiro, 1920 (Anura: Neobatrachia: Brachycephalidae), in a highland forest of southeastern Brazil. J. Nat. Hist. 2012, 46, 2497–2507. [Google Scholar] [CrossRef]
  97. Pombal, J.P., Jr.; Wistuba, E.M.; Bornschein, M.R. A new species of brachycephalid (Anura) from the Atlantic Rain Forest of Brazil. J. Herpetol. 1998, 32, 70–74. [Google Scholar] [CrossRef]
  98. Haddad, C.F.B.; Alves, A.C.R.; Clemente-Carvalho, R.B.G.; Reis, S.F. A new species of Brachycephalus from the Atlantic Rain Forest in São Paulo state, southeastern Brazil (Amphibia: Anura: Brachycephalidae). Copeia 2010, 410–420. [Google Scholar] [CrossRef]
  99. Abegg, A.D.; Ortiz, F.R.; Rocha, B.; Condes, T.H. A new record for Brachycephalus nodoterga (Amphibia, Anura Brachycephalidae) in the state of São Paulo, Brazil. Check List 2015, 11, 1769. [Google Scholar] [CrossRef]
  100. Ribeiro, R.S. Ecologia Alimentar das Quatro Espécies Dominantes da Anurofauna de Serapilheira Em Um Gradiente Altitudinal na Ilha de São Sebastião, SP. Master Thesis, Universidade Estadual Paulista “Júlio de Mesquita Filho”, Rio Claro, São Paulo, Brazil, 2006. [Google Scholar]
  101. Campos, L.A.; Silva, H.R.; Sebben, A. Morphology and development of additional bony elements in the genus Brachycephalus (Anura: Brachycephalidae). Biol. J. Linn. Soc. 2010, 99, 752–767. [Google Scholar] [CrossRef]
  102. Oliveira, E.G. História Natural de Brachycephalus pitanga no Núcleo Santa Virgínia, Parque Estadual da Serra do Mar, Estado de São Paulo. Master Thesis, Universidade Estadual Paulista “Júlio de Mesquita Filho”, Rio Claro, Brazil, 2013. [Google Scholar]
  103. Tandel, M.C.F.F.; Loibel, S.; Oliveira, E.G.; Haddad, C.F.B. Diferenciação de 3 tipos de vocalizações (cantos) na espécie Brachycephalus pitanga. Revista da Estatística da Universidade Federal de Ouro Preto 2014, 3, 374–386. [Google Scholar]
  104. Araújo, C.B.; Guerra, T.J.; Amatuzzi, M.C.O.; Campos, L.A. Advertisement and territorial calls of Brachycephalus pitanga (Anura: Brachycephalidae). Zootaxa 2012, 3302, 66–67. [Google Scholar] [CrossRef]
  105. Bornschein, M.R.; Ribeiro, L.F.; Rollo, M.M., Jr.; Confetti, A.E.; Pie, M.R. Advertisement call of Brachycephalus albolineatus (Anura: Brachycephalidae). PeerJ 2018, 6, e5273. [Google Scholar] [CrossRef]
  106. Fontoura, P.L.; Ribeiro, L.F.; Pie, M.R. Diet of Brachycephalus brunneus (Anura: Brachycephalidae) in the Atlantic Rainforest of Paraná, southern Brazil. Zoologia 2011, 28, 687–689. [Google Scholar] [CrossRef]
  107. Pie, M.R.; Ströher, P.R.; Bornschein, M.R.; Ribeiro, L.F.; Faircloth, B.C.; Mccormack, J.E. The mitochondrial genome of Brachycephalus brunneus (Anura: Brachycephalidae), with comments on the phylogenetic position of Brachycephalidae. Biochem. Syst. Ecol. 2017, 71, 26–31. [Google Scholar] [CrossRef]
  108. Monteiro, J.P.C.; Condez, T.H.; Garcia, P.C.A.; Haddad, C.F.B. The advertisement calls of two species of Brachycephalus (Anura: Brachycephalidae) from southern Atlantic Forest, Brazil. Zootaxa 2018, 4415, 183–188. [Google Scholar] [CrossRef]
  109. Wistuba, E.M. História Natural de Brachycephalus pernix Pombal, Wistuba e Bornschein, 1998 (Anura) no Morro Anhangava, Município de Quatro Barras, Estado do Paraná. Ph.D. Thesis, Universidade Federal do Paraná, Curitiba, Brazil, 1998. [Google Scholar]
  110. Pires, O.R., Jr.; Sebben, A.; Schwartz, E.F.; Morales, R.A.V.; Bloch, C., Jr.; Schwartz, C.A. Further report of the occurrence of tetrodotoxin and new analogues in the Anuran family Brachycephalidae. Toxicon 2005, 45, 73–79. [Google Scholar] [CrossRef] [PubMed]
  111. Ribeiro, L.F.; Ströher, P.R.; Firkowski, C.R.; Cini, F.A.; Bornschein, M.R.; Pie, M.R. Brachycephalus pernix (Anura: Brachycephalidae), a new host of Ophiotaenia (Eucestoda: Proteocephalidea). Herpetol. Notes 2014, 7, 291–294. [Google Scholar]
  112. Pie, M.R.; Ribeiro, L.F. A new species of Brachycephalus (Anura: Brachycephalidae) from the Quiriri mountain range of southern Brazil. PeerJ 2015, 3, e1179. [Google Scholar] [CrossRef] [PubMed]
  113. Garey, M.V.; Lima, A.M.X.; Hartmann, M.T.; Haddad, C.F.B. A new species of miniaturized toadlet, genus Brachycephalus (Anura: Brachycephalidae), from southern Brazil. Herpetologica 2012, 68, 266–271. [Google Scholar] [CrossRef]
  114. Bornschein, M.R.; Rollo, M.M., Jr.; Pie, M.R.; Confetti, A.E.; Ribeiro, L.F. Redescription of the advertisement call of Brachycephalus tridactylus (Anura: Brachycephalidae). Phyllomedusa 2019, 18, 3–12. [Google Scholar] [CrossRef]
  115. Cunha, A.K.; Oliveira, I.S.; Hartmann, M.T. Anurofauna da Colônia Castelhanos, na Área de Proteção Ambiental de Guaratuba, Serra do Mar paranaense, Brasil. Biotemas 2010, 23, 123–134. [Google Scholar]
  116. De Oliveira, A.K.C.; Oliveira, I.S.; Hartmann, M.T.; Silva, N.R.; Toledo, L.F. Amphibia, Anura, Brachycephalidae, Brachycephalus hermogenesi (Giaretta and Sawaya, 1998): New species record in the state of Paraná, southern Brazil and geographic distribution map. Check List 2011, 7, 17–18. [Google Scholar] [CrossRef]
  117. Mariotto, L.R. Anfíbios de Um Gradiente Altitudinal Em Mata Atlântica. Master Thesis, Universidade Federal do Paraná, Curitiba, Brazil, 2014. [Google Scholar]
  118. Condez, T.H.; Sawaya, R.J.; Dixo, M. Herpetofauna dos remanescentes de Mata Atlântica da região de Tapiraí e Piedade, SP, sudeste do Brasil. Biota Neotrop. 2009, 9, 157–185. [Google Scholar] [CrossRef]
  119. Verdade, V.K.; Rodrigues, M.T.; Pavan, D. Anfíbios Anuros da Região da Estação Biológica do Alto da Serra de Paranapiacaba. Patrimônio da Reserva Biológica do Alto da Serra de Paranapiacaba: A antiga Estação Biológica do Alto da Serra; Governo do Estado de São Paulo, Secretaria do Meio Ambiente: São Paulo, Brazil, 2009; pp. 579–603. [Google Scholar]
  120. Trevine, V.; Forlani, M.C.; Haddad, C.F.B.; Zaher, H. Herpetofauna of Paranapiacaba: Expanding our knowledge on a historical region in the Atlantic forest of southeastern Brazil. Zoologia 2014, 31, 126–146. [Google Scholar] [CrossRef]
  121. Collins, J.P. Amphibian decline and extinction: What we know and what we need to learn. Dis. Aquat. Organ. 2010, 92, 93–99. [Google Scholar] [CrossRef]
  122. Walter, H. Zonas de Vegetación y Clima: Breve Exposición Desde el Punto de Vista Causal y Global; Omega: Barcelona, Espanha, 1977; pp. 1–245. [Google Scholar]
  123. Walsh, R.P.D. Climate. In The Tropical Rain Forest; Richards, P.W., Ed.; Cambridge University Press: Cambridge, UK, 1979; pp. 159–205. [Google Scholar]
  124. Roderjan, C.V. O Gradiente da Floresta Ombrófila Densa no Morro Anhangava, Quatro Barras, PR—Aspectos Climáticos, Pedológicos e Fitossociológicos. Ph.D. Thesis, Universidade Federal do Paraná, Curitiba, Brazil, 1994. [Google Scholar]
  125. Pimm, S.L.; Jenkins, C.N.; Abell, R.; Brooks, T.M.; Gittleman, J.L.; Joppa, L.N.; Raven, P.H.; Roberts, C.M.; Sexton, J.O. The biodiversity of species and their rates of extinction, distribution, and protection. Science 2014, 344, 1246752. [Google Scholar] [CrossRef] [PubMed]
  126. Bornschein, M.R.; Firkowski, C.R.; Baldo, D.; Ribeiro, L.F.; Belmonte-Lopes, R.; Corrêa, L.; Morato, S.A.A.; Pie, M.R. Three new species of phytotelm-breeding Melanophryniscus from the Atlantic Rainforest of southern Brazil (Anura: Bufonidae). PLoS ONE 2015, 10, e0142791. [Google Scholar] [CrossRef] [PubMed]
  127. Nadaline, M.J.; Ribeiro, L.F.; Teixeira, L.; Vannuchi, F.S.; Bornschein, M.R. New record of Melanophryniscus biancae Bornschein, Baldo, Pie, Firkowski, Ribeiro & Corrêa, 2015 (Anura: Bufonidae) from Paraná, Brazil, with comments on its phytotelm-breeding ecology. Check List, in press.
Figure 1. Flow chart indicating the approach to creating polygons of the extent of occurrence to compare the results with IUCN’s species extinction risk classification criteria [48].
Figure 1. Flow chart indicating the approach to creating polygons of the extent of occurrence to compare the results with IUCN’s species extinction risk classification criteria [48].
Diversity 11 00150 g001
Figure 2. Status of conservation for the 36 species of Brachycephalus proposed in this study according to the IUCN [48] criteria and categories proposed by IUCN [31,32,33,34,35,36,37,38,39,40,41]. Abbreviation: CR = Critically Endangered; EN = Endangered; VU = Vulnerable; LC = Least Concern; DD = Data Deficient.
Figure 2. Status of conservation for the 36 species of Brachycephalus proposed in this study according to the IUCN [48] criteria and categories proposed by IUCN [31,32,33,34,35,36,37,38,39,40,41]. Abbreviation: CR = Critically Endangered; EN = Endangered; VU = Vulnerable; LC = Least Concern; DD = Data Deficient.
Diversity 11 00150 g002
Table 1. Locality records of Brachycephalus.
Table 1. Locality records of Brachycephalus.
SpeciesGroupLocality and StateAltitude 1Source
B. didactylusdidactylusMonumento Natural Serra das Torres (21°00’04” S, 41°13’17” W), municipality of Atílio Vivácqua, Espírito Santo600–900?[51] as B. didactylus; [52] as B. didactylus
B. didactylusdidactylusFazenda Santa Bárbara (22°25’17” S, 42°35’01” W), Parque Estadual dos Três Picos, municipality of Cachoeiras de Macacu, Rio de Janeiro500–800[53] as B. didactylus; [54] as B. didactylus
B. didactylusdidactylusReserva Ecológica de Guapiaçu (22°24’00” S, 42°44’00” W), municipality of Cachoeiras de Macacu, Rio de Janeiro300–520[55] as B. didactylus]
B. didactylusdidactylusReserva Ecológica Rio das Pedras (22°59’00” S, 44°06’45” W), municipality of Mangaratiba, Rio de Janeiro200–1110[23] as B. didactylus; [54] as B. didactylus; [56] as B. didactylus; [57] as B. didactylus
B. didactylusdidactylusSacra Família do Tinguá (22°29’11” S, 43°36’18” W), municipality of Engenheiro Paulo de Frontin, Rio de Janeiro600[17] as B. didactylus; [27] as B. didactylus; [58] as B. didactylus; [59] as B. didactylus; [60] as B. didactylus; [61] as B. didactylus; [62] as B. didactylus; [63] as B. didactylus; [64] as B. didactylus; [65] as B. didactylus; [66] as B. didactylus
B. didactylusdidactylusTheodoro de Oliveira (first position: 22°22’11” S, 42°33’25” W), Parque Estadual dos Três Picos, municipality of Nova Friburgo, Rio de Janeiro1100–1400?[23] as B. didactylus; [67] as B. didactylus
B. didactylusdidactylusTinguá (22°35’51” S, 43°24’54” W), municipality of Nova Iguaçu, Rio de Janeiro35[17] as B. didactylus
B. didactylusdidactylusVila Dois Rios (23°11’01” S, 44°12’23” W), Ilha Grande, municipality of Angra dos Reis, Rio de Janeiro220–240[23] as B. didactylus; [68] as B. didactylus; [69] as B. didactylus; [70] as B. didactylus
B. hermogenesididactylusCorcovado (23°28’20” S, 45°11’41” W), municipality of Ubatuba, São Paulo30–250This study, [18] as B. hermogenesi; [23] as B. hermogenesi; [25] as B. hermogenesi collected at Picinguaba; [27] as B. hermogenesi; [63] as B. hermogenesi
B. hermogenesididactylusEstação Biológica de Boracéia (23°39’10” S, 45°53’05” W), municipality of Salesópolis, São Paulo825–900[23] as B. hermogenesi; [27] as B. hermogenesi; [63] as B. hermogenesi; [71] as B. hermogenesi; [72] as B. hermogenesi
B. hermogenesididactylusFazenda Capricórnio (23°23’27” S, 45°04’26” W), municipality of Ubatuba, São Paulo60[18] as B. hermogenesi; [23] as B. hermogenesi; [27] as B. hermogenesi; [63] as B. hermogenesi; [72] as B. hermogenesi
B. hermogenesididactylusMorro Cuscuzeiro (23°17’50”S, 44°47’21” W), on the border of municipalities of Paraty, Rio de Janeiro, and Ubatuba, São Paulo730–1090This study
B. hermogenesididactylusMorro do Corcovado (23°27’06” S, 45°12’03” W), Parque Estadual da Serra do Mar, municipality of Ubatuba, São Paulo250–1060This study
B. hermogenesididactylusMunicipality of Paraibuna (c. 23°23’34” S, 45°39’42” W), São Paulo?[72] as B. hermogenesi
B. hermogenesididactylusNúcleo Cunha (23°15’48”S, 45°02’39”W), Parque Estadual da Serra do Mar, municipality of Cunha, São Paulo1045–1140This study
B. hermogenesididactylusNúcleo Picinguaba (23°22’21”S, 44°49’53”W), Parque Estadual da Serra do Mar, municipality of Ubatuba, São Paulo0–700[18] as B. hermogenesi; [23] as B. hermogenesi]; [27] as B. hermogenesi; [29] as B. hermogenesi; [63] as B. hermogenesi; [64] as B. hermogenesi; [71] as B. hermogenesi; [72] as B. hermogenesi
B. hermogenesididactylusSertão da Cutia (not located), municipality of Ubatuba, São Paulo?[72] as B. hermogenesi
B. hermogenesididactylusTrilha do Corisco (23°16’38” S, 44°46’39” W), municipality of Paraty, Rio de Janeiro350–725This study
B. hermogenesididactylusTrilha do Ipiranga 50 m from the Rio Ipiranga (23°20’41” S, 45°08’21” W), Núcleo Santa Virgínia, Parque Estadual da Serra do Mar, municipality of São Luiz do Paraitinga, São Paulo920–940This study
B. pulexdidactylusSerra Bonita (15°23’28” S, 39°33’59” W), municipality of Camacan, Bahia800–930[20] as B. pulex
B. sulfuratusdidactylusBase of the Serra Água Limpa (24°28’52” S, 48°47’12” W), municipality of Apiaí, São Paulo920[23] as Brachycephalus sp. 1; [25] as B. sulfuratus; [28] without species identification; [50] as B. sulfuratus; [73] without species identification; [74] as B. sulfuratus
B. sulfuratusdidactylusBiquinha (24°17’43” S, 47°36’26” W), municipality of Juquiá, São Paulo40This study
B. sulfuratusdidactylusBraço do Norte (26°07’29” S, 48°43’48” W), municipality of Itapoá, Santa Catarina240[75] as B. sulfuratus
B. sulfuratusdidactylusCaratuval, near the Parque Estadual das Lauráceas (24°51’17” S, 48°43’43” W), municipality of Adrianópolis, Paraná900[23] as Brachycephalus sp. 1; [25] as B. sulfuratus; [27] as Brachycephalus sp. nov. 1; [28] without species identification; [50] as B. sulfuratus; [73] without species identification; [74] as B. sulfuratus
B. sulfuratusdidactylusCaratuval, Parque Estadual das Lauráceas (24°51’14” S, 48°42’01” W), municipality of Adrianópolis, Paraná890[23] as Brachycephalus sp. 1; [27] as Brachycephalus sp. nov. 1
B. sulfuratusdidactylusCastelo dos Bugres (26°13’47” S, 49°03’20” W), municipality of Joinville, Paraná790–860[23] as Brachycephalus sp. 1; [27] as Brachycephalus sp. nov. 1; [72] as B. sulfuratus; [75] as B. sulfuratus
B. sulfuratusdidactylusCentro de Estudos e Pesquisas Ambientais da Univille (26°13’39” S, 48°41’31” W), Vila da Glória, Distrito do Saí, municipality of São Francisco do Sul, Santa Catarina125[72] as B. sulfuratus
B. sulfuratusdidactylusCorvo (25°20’17” S, 48°54’56” W), municipality of Quatro Barras, Paraná930[23] as Brachycephalus sp. 1; [25] as B. sulfuratus; [27] as Brachycephalus sp. nov. 1; [28] without species identification; [29] as B. sulfuratus; [50] as B. sulfuratus; [73] without species identification; [74] as B. sulfuratus
B. sulfuratusdidactylusEstância Hidroclimática Recreio da Serra (25°27’14” S, 49°00’28” W), Serra da Baitaca, municipality of Piraquara, Paraná1150–1205This study
B. sulfuratusdidactylusFazenda Thalia (25°30’58” S, 49°40’12” W), municipality of Balsa Nova, Paraná1025[23] as Brachycephalus sp. 1; [25] as B. sulfuratus; [27] as Brachycephalus sp. nov. 1; [28] without species identification; [50] as B. sulfuratus; [73] without species identification; [74] as B. sulfuratus
B. sulfuratusdidactylusnear the Jurupará dam (23°56’30” S, 47°23’45” W), municipality of Piedade, São Paulo690[25] as B. sulfuratus
B. sulfuratusdidactylusMananciais da Serra (25°29’32” S, 48°59’33” W), municipality of Piraquara, Paraná970–1050[23] as Brachycephalus sp. 1; [25] as B. sulfuratus; [27] as Brachycephalus sp. nov. 1; [50] as B. sulfuratus; [74] as B. sulfuratus
B. sulfuratusdidactylusMorro Anhangava (25°22’51” S, 49°01’26” W), municipality of Quatro Barras, Paraná915[72] as B. sulfuratus; [75] as B. sulfuratus
B. sulfuratusdidactylusMorro do Cantagalo (26°10’31” S, 48°42’44” W), Vila da Glória, Distrito do Saí, municipality of São Francisco do Sul, Santa Catarina160[72] as B. sulfuratus
B. sulfuratusdidactylusMorro do Garrafão (26°28’23” S, 49°15’57” W), municipality of Corupá, Santa Catarina500–530[25] as B. sulfuratus; [76] as B. sulfuratus
B. sulfuratusdidactylusMorro Garuva (26°02’29” S, 48°53’14” W), municipality of Garuva, Santa Catarina215–495This study
B. sulfuratusdidactylusNúcleo Itutinga-Pilões (23°54’17” S, 46°29’22” W), Parque Estadual da Serra do Mar, municipality of Cubatão, São Paulo55This study
B. sulfuratusdidactylusParque Estadual da Ilha do Cardoso (25°06’53” S, 47°55’40” W), municipality of Cananéia, São Paulo385[63] as possibly B. hermogenesi; [72] as B. sulfuratus
B. sulfuratusdidactylusParque Estadual Intervales (24°16’33” S, 48°25’04” W), municipality of Iporanga, São Paulo820This study
B. sulfuratusdidactylusRecanto das Hortências (25°33’24” S, 48°59’38” W), municipality of São José dos Pinhais, Paraná975[23] as Brachycephalus sp. 1; [25] as B. sulfuratus; [50] as B. sulfuratus; [74] as B. sulfuratus
B. sulfuratusdidactylusReserva Particular do Patrimônio Natural Salto Morato (25°09’14” S, 48°18’06” W), municipality of Guaraqueçaba, Paraná40–880[23] as Brachycephalus sp. 1; [77] as B. hermogenesi; [78] as B. hermogenesi; [79] as B. hermogenesi
B. sulfuratusdidactylusSalto do Inferno (25°00’02” S, 48°37’07” W), Rio Capivari, municipality of Bocaiúva do Sul, Paraná610[25] as B. sulfuratus; [50] as B. sulfuratus; [74] as B. sulfuratus
B. sulfuratusdidactylusSerra do Guaraú (24°47’12” S, 48°07’11” W), on the border of the municipalities of Cajati and Jacupiranga, São Paulo680–835This study
B. sulfuratusdidactylusSerra do Pico (26°08’31” S, 48°57’19” W), municipality of Joinville, Santa Catarina340–720This study
B. sulfuratusdidactylusTorre Embratel (24°52’46” S, 48°15’27” W), municipality of Cajati, São Paulo960–990This study
B. sulfuratusdidactylusTruticultura (26°01’33” S, 48°52’02” W), municipality of Garuva, Paraná90[23] as Brachycephalus sp. 1; [27] as Brachycephalus sp. nov. 1
B. alipioiephippiumFazenda Aoki or Fazenda dos Japoneses (20°28’24” S, 41°00’36” W), boundary of the municipalities of Vargem Alta and Domingos Martins, Espírito Santo1070–1100[27] as B. alipioi; [64] as B. alipioi; [66] as B. alipioi; [80] as B. alipioi; [81] as B. alipioi; [82] as B. alipioi
B. alipioiephippiumForno Grande (20°31’41” S, 41°06’51” W), Parque Estadual de Forno Grande, municipality of Castelo, Espírito Santo1430?[27] as B. alipioi
B. alipioiephippiumAlto Castelinho (20°30’34” S, 41°00’33” W), municipality of Vargem Alta, Espírito Santo1100This study, [25] as B. alipioi
B. bufonoidesephippiumSerra de Macaé (22°18’02” S, 42°18’20” W), municipality of Nova Friburgo, Rio de Janeiro1100?[30] as B. bufonoides; [66] as B. bufonoides; [83] as B. ephippium bufonoides
B. crispusephippiumBacia B, Núcleo Cunha, Parque Estadual da Serra do Mar (23°15’15” S, 45°01’58” W), municipality of Cunha, São Paulo800–1190This study, [84] as B. crispus
B. darksideephippiumMata do Pai Inácio (20°46′44″ S, 42°29′10″ W), Parque Estadual da Serra do Brigadeiro, municipality Miradouro, Minas Gerais1340[66] as B. ephippium; [85] as B. ephippium; [86] as B. darkside
B. darksideephippiumTrilha do Cruzeiro (20°52′41″ S, 42°31′15″ W), Parque Estadual da Serra do Brigadeiro, boundary of the municipalities of Ervália and Muriaé, Minas Gerais1265–1500[86] as B. darkside
B. ephippiumephippiumCondomínio Ermida (23°14’13” S, 46°58’52” W), Serra do Japi, municipality of Jundiaí, São Paulo1225[27] as B. ephippium
B. ephippiumephippiumHotel Fazenda Pé da Serra (22°51’56” S, 45°31’40” W), municipality of Pindamonhangaba, São Paulo700[27] as B. ephippium
B. ephippiumephippiumLago Azul (22°27’23” S, 44°36’34” W), Parque Nacional do Itatiaia, municipality of Itatiaia, Rio de Janeiro750[27] as B. ephippium
B. ephippiumephippiumMaromba (22°25’43” S, 44°37’11” W), Parque Nacional do Itatiaia, municipality of Itatiaia, Rio de Janeiro1125[27] as B. ephippium
B. ephippiumephippiumMonteiro Lobato (22°57’07” S, 45°50’20” W), municipality of Monteiro Lobato, São Paulo700[66] as B. ephippium
B. ephippiumephippiumObservatório de Capricórnio (22°53’54” S, 46°49’01” W), Serra das Cabras, Joaquim Egídio District, boundary of the municipalities of Campinas and Morungaba, São Paulo1085[19] as B. ephippium]; [27] as B. ephippium; [66] as B. ephippium; [87] as B. ephippium
B. ephippiumephippiumParque Municipal de Itapetinga (Grota Funda) (23°11’07” S, 46°31’47” W), municipality of Atibaia, São Paulo900–1250[27] as B. ephippium; [64] as B. ephippium; [81] as B. ephippium; [88] as B. ephippium; [89] as B. ephippium
B. ephippiumephippiumReserva Biológica da Serra do Japi (23°17’07” S, 47°00’05” W), Serra do Japi, boundary of the municipalities of Jundiaí and Cabreúva, São Paulo1000[27] as B. ephippium; [64] as B. ephippium; [66] as B. ephippium; [90] as B. ephippium
B. ephippiumephippiumReserva Ecológica do Trabiju (22°48’01” S, 45°32’03” W), Trabiju, municipality of Pindamonhangaba, São Paulo1000?[66] as B. ephippium
B. ephippiumephippiumReserva Pedra Branca (22°56’22” S, 45°41’04” W), municipality of Tremembé, São Paulo890?[66] as B. ephippium
B. ephippiumephippiumSanto Antônio do Pinhal (22°49’28” S, 45°40’20” W), municipality of Santo Antônio do Pinhal, São Paulo1080[66] as B. ephippium
B. ephippiumephippiumSão Francisco Xavier (22°53’44” S, 45°58’04” W), municipality of São José dos Campos, São Paulo1000[27] as B. ephippium; [66] as B. ephippium; [91] as B. ephippium; [92] as B. ephippium
B. ephippiumephippiumSerra Negra (21°57’28” S, 43°47’20” W), municipality of Santa Bárbara do Monte Verde, Minas Gerais?[23] as B. ephippium; [65] as BMV MG2
B. ephippiumephippiumSerra da Concórdia (22°20’30” S, 43°44’04” W), Parque Estadual Serra da Concórdia, Barão de Juparanã, municipality of Valença, Rio de Janeiro900?[66] as B. ephippium
B. ephippiumephippiumAlto do Soberbo (22°27’15” S, 42°59’21” W), municipality of Teresópolis, Rio de Janeiro1250[66] as B. ephippium
B. ephippiumephippiumComary (22°27’22” S, 42°58’24” W), municipality of Teresópolis, Rio de Janeiro990[66] as B. ephippium
B. ephippiumephippiumFloresta dos Macacos (22°58’15” S, 43°15’24” W), municipality of Rio de Janeiro, Rio de Janeiro450?[66] as B. ephippium
B. ephippiumephippiumGarrafão (22°28’04” S, 43°01’52” W), municipality of Guapimirim, Rio de Janeiro1785?[66] as B. ephippium
B. ephippiumephippiumPedra Branca (22°55’55” S, 43°28’23” W), Serra da Pedra Branca, municipality of Rio de Janeiro, Rio de Janeiro1000[58] as B. ephippium; [66] as B. ephippium
B. ephippiumephippiumRepresa do Rio Grande (22°55’58” S, 43°26’36” W), Parque Estadual da Pedra Branca, municipality of Rio de Janeiro, Rio de Janeiro150?[27] as B. ephippium; [66] as B. ephippium
B. ephippiumephippiumReserva Ecológica Rio das Pedras (22°59’00” S, 44°06’45” W), municipality of Mangaratiba, Rio de Janeiro200–1110[56] as B. ephippium
B. ephippiumephippiumRiacho Beija-flor (22°27’04” S, 43°00’04” W), Parque Nacional da Serra dos Órgãos, municipality of Teresópolis, Rio de Janeiro1195[27] as B. ephippium
B. ephippiumephippiumRocio District (22°28’23” S, 43°14’38” W), municipality of Petrópolis, Rio de Janeiro950[27] as B. ephippium
B. ephippiumephippiumSerra do Tinguá (22°35’31” S, 43°28’16” W), municipality of Nova Iguaçu, Rio de Janeiro950?[66] as B. ephippium
B. ephippiumephippiumVale da Revolta (22°26’17” S, 42°56’19” W), municipality of Teresópolis, Rio de Janeiro1035[66] as B. ephippium
B. ephippiumephippiumVarginha (22°24’34” S, 42°52’11” W), municipality of Teresópolis, Rio de Janeiro825?[66] as B. ephippium
B. ephippiumephippiumBonito (22°42’51” S, 44°34’39” W), Serra da Bocaina, municipality of São José do Barreiro, São Paulo1660?[66] as B. ephippium
B. ephippiumephippiumEstação Ecológica de Bananal (22°48’05” S, 44°22’12” W), Serra da Bocaina, municipality of Bananal, São Paulo1200?[93] as B. ephippium
B. ephippiumephippiumLídice District (22°50’01” S, 44°11’32” W), municipality of Rio Claro, Rio de Janeiro650?[58] as B. ephippium; [66] as B. ephippium
B. ephippiumephippiumPedra Branca (23°10’38” S, 44°47’19” W), Serra da Bocaina, municipality of Parati, Rio de Janeiro630?[58] as B. ephippium; [66] as B. ephippium
B. ephippiumephippiumReserva Florestal de Morro Grande (23°42’08” S, 46°58’22” W), municipality of Cotia, São Paulo990?[94] as B. ephippium
B. garbeanusephippiumAlto Caledônia (22°20’10” S, 42°33’20” W), municipality of Nova Friburgo, Rio de Janeiro1070?[66] as B. garbeanus
B. garbeanusephippiumBaixo Caledônia (22°21’33” S, 42°34’12” W), municipality of Nova Friburgo, Rio de Janeiro1600–1900[66] as B. garbeanus; [67] as B. garbeanus; [95] as B. garbeanus; [96] as B. garbeanus
B. garbeanusephippiumMacaé de Cima (22°21’37” S, 42°17’50” W), municipality of Nova Friburgo, Rio de Janeiro1130[27] as B. garbeanus; [64] as B. ephippium; [66] as B. garbeanus; [81] as B. garbeanus; [91] as B. ephippium; [92] as B. ephippium
B. garbeanusephippiumMorro São João (22°22’47” S, 42°30’34” W), municipality of Nova Friburgo, Rio de Janeiro1550?[66] as B. garbeanus
B. garbeanusephippiumSerra de Macaé (22°18’02” S, 42°18’20” W), municipality of Nova Friburgo, Rio de Janeiro1100?[30] as B. garbeanus; [66] as B. garbeanus; [83] as B. ephippium garbeanus
B. garbeanusephippiumSerra Nevada (22°21’46” S, 42°32’48” W), municipality of Nova Friburgo, Rio de Janeiro1190[66] as B. garbeanus
B. garbeanusephippiumTheodoro de Oliveira (second position: 22°21’48” S, 42°33’13” W), Parque Estadual dos Três Picos, municipality of Nova Friburgo, Rio de Janeiro1400[66] as B. garbeanus; [67] as B. garbeanus; [95] as B. garbeanus
B. guaraniephippiumMorro Prumirim (23°20’50” S, 45°01’37” W), municipality of Ubatuba, São Paulo500–900[82] as B. guarani; [84] as B. guarani; [88] as Brachycephalus sp.
B. margaritatusephippiumCastelo Country Club (22°32’21” S, 43°13’08” W), municipality of Petrópolis, Rio de Janeiro980[66] as B. margaritatus
B. margaritatusephippiumCastelo Montebello (22°24’24” S, 42°58’06” W), municipality of Teresópolis, Rio de Janeiro920[66] as B. margaritatus
B. margaritatusephippiumIndependência (22°32’58” S, 43°12’27” W), municipality of Petrópolis, Rio de Janeiro860[66] as B. margaritatus
B. margaritatusephippiumMorro Azul (22°28’34” S, 43°34’40” W), municipality of Engenheiro Paulo de Frontin, Rio de Janeiro620[65] as BPF RJ2; [66] as B. margaritatus
B. margaritatusephippiumQuitandinha (22°31’47” S, 43°12’26” W), municipality of Petrópolis, Rio de Janeiro925[66] as B. margaritatus
B. margaritatusephippiumSacra Família do Tinguá (22°29’11” S, 43°36’18” W), municipality of Engenheiro Paulo de Frontin, Rio de Janeiro600[17] as B. ephippium; [27] as B. ephippium; [58] as Brachycephalus cf. ephippium; [66] as B. margaritatus
B. nodotergaephippiumEstação Biológica de Boracéia (second position: 23°38’00”S, 45°52’00”W), municipality of Salesópolis, São Paulo945[27] as B. nodoterga; [30] as B. nodoterga; [58] as B. nodoterga; [59] as B. nodoterga; [60] as B. nodoterga; [61] as B. nodoterga; [66] as B. nodoterga; [97] as B. nodoterga; [98] as B. nodoterga; [99] as B. nodoterga
B. nodotergaephippiumFazenda Paiva Ramos (23°28’21” S, 46°47’25” W), municipality of Osasco, São Paulo820[99] as B. nodoterga
B. nodotergaephippiumPico do Ramalho (23°51’42” S, 45°21’28” W), Ilha de São Sebastião, municipality of Ilhabela, São Paulo700–900[27] as B. nodoterga; [66] as B. nodoterga; [99] as B. nodoterga; [100] as Brachycephalus sp. aff. nodoterga
B. nodotergaephippiumSantana de Parnaíba (23°26’19” S, 46°56’06” W), municipality of Santana de Parnaíba, São Paulo730[99] as B. nodoterga
B. nodotergaephippiumSerra da Cantareira (23°27’13” S, 46°38’11” W), Parque Estadual da Cantareira, municipality of São Paulo, São Paulo850?[30] as B. nodoterga; [59] as B. nodoterga; [60] as B. nodoterga; [61] as B. nodoterga; [64] as B. nodoterga; [66] as B. nodoterga; [81] as B. nodoterga; [82] as B. nodoterga, [83] as B. ephippium nodoterga; [84] as B. nodoterga, [98] as B. nodoterga; [99] as B. nodoterga
B. pitangaephippiumFazenda Capricórnio (23°22’36” S, 45°04’07” W), municipality of Ubatuba, São Paulo450?[27] as B. pitanga; [61] as B. pitanga; [65] as B. pitanga; [101] as Brachycephalus sp. 2
B. pitangaephippiumNúcleo Santa Virgínia (23°19’23” S, 45°05’19” W), Parque Estadual da Serra do Mar, municipality of São Luis do Paraitinga, São Paulo980–1140[102] as B. pitanga; [103] as B. pitanga
B. pitangaephippiumSP 125—municipality of São Luís do Paraitinga (23°22’57” S, 45°09’59” W), São Paulo935–950[23] as B. pitanga
B. pitangaephippiumTrilha do Ipiranga 50 m from the Rio Ipiranga (23°20’39” S, 45°08’16” W), Núcleo Santa Virgínia, Parque Estadual da Serra do Mar, municipality of São Luis do Paraitinga, São Paulo900–960[27] as B. pitanga; [61] as B. pitanga; [64] as B. pitanga; [81] as B. pitanga; [102] as B. pitanga; [104] as B. pitanga
B. tobyephippiumMorro do Corcovado (23°27’22” S, 45°11’53” W), Parque Estadual da Serra do Mar, municipality of Ubatuba, São Paulo750–1060This study, [27] as B. toby; [81] as B. toby; [82] as B. toby; [84] as B. toby; [98] as B. toby
B. vertebralisephippiumMorro Cuscuzeiro (23°17’51” S, 44°47’20” W), on the border of municipalities of Paraty, Rio de Janeiro, and Ubatuba, Sao Paulo760–1110This study, [27] as B. vertebralis; [81] as B. vertebralis; [84] as B. vertebralis
B. vertebralisephippiumPedra Branca (23°10’38” S, 44°47’19” W), Serra da Bocaina, municipality of Parati, Rio de Janeiro630?[27] as B. vertebralis; [30] as B. vertebralis; [58] as. B. vertebralis; [64] as B. vertebralis
B. actaeuspernixBraço do Norte (26°07′29″ S, 48°43′48″ W), municipality of Itapoá, Santa Catarina240[75] as B. actaeus
B. actaeuspernixCentro de Estudos e Pesquisas Ambientais da Univille (CEPA) (26°13’39” S; 48°41’31” W), Vila da Glória, Distrito do Saí, municipality of São Francisco do Sul, Santa Catarina120[75] as B. actaeus
B. actaeuspernixEstrada do Saí (26°12′06″ S; 48°41′37″ W), Distrito do Saí, municipality of São Francisco do Sul, Santa Catarina100[75] as B. actaeus
B. actaeuspernixFazenda Morro Grande (26°17′47″ S; 48°37′10″ W), Morro Grande, Ilha de São Francisco, municipality of São Francisco do Sul, Santa Catarina60[75] as B. actaeus
B. actaeuspernixFazenda Palmito Juriti (26°08′09″ S; 48°43′54″ W), municipality of Itapoá, Santa Catarina100–170[75] as B. actaeus
B. actaeuspernixSerra da Palha (26°17′50″ S; 48°40′28″ W), Laranjeiras, Ilha de São Francisco, municipality of São Francisco do Sul, Santa Catarina20–90[75] as B. actaeus
B. actaeuspernixSerra da Tiririca (26°07′42″ S, 48°44′32″ W), municipality of Itapoá, Santa Catarina170–530This study
B. albolineatuspernixMorro Azul (26°45′52″ S, 49°12′20″ W), on the border between the municipalities of Pomerode and Rio dos Cedros, Santa Catarina725–740This study
B. albolineatuspernixMorro Boa Vista (26°30’58” S, 49°03’14” W), on the border between the municipalities of Jaraguá do Sul and Massaranduba, Santa Catarina790–835[74] as B. albolineatus; [105] as B. albolineatus
B. albolineatuspernixMorro do Garrafão (26°30’58” S, 49°03’14” W), municipality of Corupá, Santa Catarina500–530[76] as B. albolineatus
B. albolineatuspernixMorro do Schmidt (26°39′55″ S, 49°12′55″ W), municipality of Pomerode, Santa Catarina810–870This study
B. auroguttatuspernixPedra da Tartaruga (26°00’21”S, 48°55’25”W), municipality of Garuva, Santa Catarina1070–1100[23] as B. auroguttatus; [26] as B. auroguttatus; [28] as B. auroguttatus; [29] as B. auroguttatus; [73] without species identification
B. boticariopernixMorro do Cachorro (26°46’42” S, 49°01’57” W), boundary of the municipalities of Blumenau, Gaspar, and Luiz Alves, Santa Catarina685–795[23] as B. boticario; [26] as B. boticario; [28] as B. boticario; [29] as B. boticario; [73] without species identification
B. brunneuspernixAbrigo 1 (25°13’29” S, 48°51’17” W), municipality of Campina Grande do Sul, Paraná1440–1640This study, [28] as not identified; [29] as B. brunneus
B. brunneuspernixCamapuã (25°15’59” S, 48°50’16” W), Serra dos Órgãos, boundary of the municipalities of Campina Grande do Sul and Antonina, Paraná1595[27] as B. brunneus; [28] as B. brunneus]; [29] as B. brunneus; [73] without species identification; [106] as B. brunneus
B. brunneuspernixCaranguejeira (25°20’27” S, 48°54’31” W), Serra da Graciosa, municipality of Quatro Barras, Paraná1095–1110[23] as B. brunneus; [73] without species identification
B. brunneuspernixCaratuva (25°14’33” S, 48°50’04” W), Serra dos Órgãos, municipality of Campina Grande do Sul, Paraná1300–1770[27] as B. brunneus; [28] as B. brunneus; [29] as B. brunneus, [59] as B. brunneus; [64] as B. brunneus; [65] as B. brunneus; [66] as B. brunneus; including Pico Paraná; [73] without species identification; including Pico Paraná; [81] as B. brunneus; [106] as B. brunneus; [107] as B. brunneus
B. brunneuspernixGetúlio (25°14’18” S, 48°50’13” W), Serra dos Órgãos, municipality of Campina Grande do Sul, Paraná1310–1490[23] as B. brunneus; [27] as B. brunneus
B. brunneuspernixMãe Catira (25°20’51” S, 48°54’25” W), Serra da Graciosa, municipality of Quatro Barras, Paraná1135–1405This study, [27] as Brachycephalus sp. nov. 2; [28] as not identified; [73] without species identification
B. coloratuspernixEstância Hidroclimática Recreio da Serra (25°27’14” S, 49°00’27” W), Serra da Baitaca, municipality of Piraquara, Paraná1145–1230[50] as B. coloratus
B. curupirapernixMorro do Canal (25°30’55” S, 48°58’56” W), municipality of Piraquara, Paraná1320This study, [23] as Brachycephalus sp. 4; [28] as not identified]; [73] without species identification
B. curupirapernixMorro do Vigia (25°30’33” S, 48°58’58” W), municipality of Piraquara, Paraná1250[23] as Brachycephalus sp. 4; [27] as Brachycephalus sp. nov. 3; [28] as not identified; [29] as B. curupira; [73] without species identification
B. curupirapernixSerra do Salto (25°42’07” S, 49°03’44” W), Malhada District, municipality of São José dos Pinhais, Paraná1095–1160[23] as Brachycephalus sp. 6; [27] as Brachycephalus sp. 2; [28] as not identified; [29] as B. curupira; [50] as B. curupira, [73] without species identification
B. ferruginuspernixOlimpo (25°27’03” S, 48°54’59” W), Serra do Marumbi, municipality of Morretes, Paraná965–1470[27] as B. ferruginus; [28] as B. ferruginus; [29] as B. ferruginus, [60] as B. ferruginus; [64] as B. ferruginus; [66] as B. ferruginus; [73] without species identification; [81] as B. ferruginus
B. fuscolineatuspernixMorro Braço da Onça (26°44’58” S, 48°55’41” W), municipality of Luiz Alves, Santa Catarina525–530[24] as B. fuscolineatus
B. fuscolineatuspernixMorro do Baú (26°47’58” S, 48°55’47” W), municipality of Ilhota, Santa Catarina640–790[26] as B. fuscolineatus; [27] as Brachycephalus sp. nov. 9; [28] as B. fuscolineatus; [29] as B. fuscolineatus; [73] without species identification
B. izecksohnipernixTorre da Prata, Serra da Prata (25°37’25” S, 48°41’31” W), boundary of the municipalities of Morretes, Paranaguá, and Guaratuba, Paraná980–1340[27] as B. izecksohni; [28] as B. izecksohni; [29] as B. izecksohni; [59] as B. izecksohni; [64] as B. izecksohni; [66] as B. izecksohni; [73] without species identification; [81] as B. izecksohni
B. leoparduspernixMorro dos Perdidos (25°53’22” S, 48°57’22” W), municipality of Guaratuba, Paraná1340–1420[26] as B. leopardus; [27] as Brachycephalus sp. nov. 4; [28] as B. leopardus; [73] without species identification
B. leoparduspernixSerra do Araçatuba (25°54’07” S, 48°59’47” W), municipality of Tijucas do Sul, Paraná1640–1645[26] as B. leopardus; [27] as Brachycephalus sp. nov. 4; [28] as B. leopardus; [73] without species identification
B. mariaeterezaepernixReserva Particular do Patrimônio Natural Caetezal, top of the Serra Queimada (26°06’51” S, 49°03’45” W), municipality of Joinville, Santa Catarina1265–1270[26] as B. mariaeterezae; [27] as Brachycephalus sp. nov. 6; [28] as B. mariaeterezae; [29] as B. mariaeterezae; [73] without species identification
B. mirissimuspernixMorro Santo Anjo (26°37’41” S, 48°55’50” W), municipality of Massaranduba, Santa Catarina470–540[25] as B. mirissimus
B. olivaceuspernixBase of the Serra Queimada (26°04’57” S, 49°03’59” W), municipality of Joinville, Santa Catarina985[17] as Brachycephalus sp. nov. 7; [26] as B. olivaceus
B. olivaceuspernixCastelo dos Bugres (second position: 26°13’59”S, 49°03’13”W), municipality of Joinville, Santa Catarina800–835[26] as B. olivaceus; [27] as Brachycephalus sp. nov. 7; [28] as B. olivaceus; [73] without species identification; [108] as B. olivaceus
B. olivaceuspernixMorro do Boi (26°24’42” S, 49°12’59” W), municipality of Corupá, Santa Catarina650–920[23] as B. olivaceus; [27] as Brachycephalus sp. 3; [29] as B. olivaceus
B. olivaceuspernixPico Jurapê (26°16′27″ S, 49°00′13″ W), municipality of Joinville, Santa Catarina650–780This study
B. pernixpernixAnhangava (25°23’19” S, 49°00’15” W), Serra da Baitaca, municipality of Quatro Barras, Paraná1135–1405[27] as B. pernix; [28] as B. pernix; [29] as B. pernix; [62] as B. pernix; [64] as B. pernix; [65] as B. pernix; [66] as B. pernix; [73] without species identification; [81] as B. pernix; [97] as B. pernix; [101] as B. pernix; [109] as B. pernix; [110] as B. pernix; [111] as B. pernix
B. pombalipernixMorro dos Padres (25°36’40” S, 48°51’22” W), Serra da Igreja, municipality of Morretes, Paraná1060–1300[27] as B. pombali; [28] as B. pombali; [29] as B. pombali; [60] as B. pombali; [64] as B. pombali; [73] without species identification; [81] as B. pombali
B. pombalipernixtrail to Morro dos Padres (25°35’58” S, 48°51’57” W), municipality of Morretes, Paraná845–1060[27] as B. pombali
B. quiririensispernixSerra do Quiriri (26°01’17” S, 48°59’47” W), municipality of Campo Alegre, Santa Catarina1240–1270[23] as B. quiririensis; [27] as Brachycephalus sp. nov. 5; [28] as B. quiririensis; [29] as B. quiririensis; [73] without species identification; [112] as B. quiririensis
B. quiririensispernixSerra do Quiriri (first position: 26°01’42” S, 48°57’11” W; second position: 26°01’32” S, 48°58’24” W), municipality of Garuva, Santa Catarina1320–1380[27] as Brachycephalus sp. nov. 5; [108] as B. quiririensis
B. tridactyluspernixSerra do Morato (25°08’09” S, 48°17’59” W), Reserva Natural Salto Morato, municipality of Guaraqueçaba, Paraná805–910[23] as B. tridactylus; [28] as B. tridactylus; [113] as B. tridactylus; [114] as B. tridactylus
B. verrucosuspernixMorro da Tromba (26°12’44” S, 48°57’29” W), municipality of Joinville, Santa Catarina455–945[23] as B. verrucosus; [26] as B. verrucosus; [27], as Brachycephalus sp. nov. 8; [28] as B. verrucosus; [29] as B. verrucosus; [73] without species identification
B. atelopoide?Piquete, São Paulo?[30] as B. atelopoide; [83] as B. ephippium atelopoide]
Brachycephalus sp. (cf. B. sulfuratus)didactylusAlto Quiriri (26°05’34” S, 48°59’41” W), municipality of Garuva, Santa Catarina240[23] as Brachycephalus sp. 1; [27] as Brachycephalus sp. nov. 1
Brachycephalus sp. (cf. B. sulfuratus)didactylusColônia Castelhanos (25°47’58” S, 48°54’40” W), municipality of Guaratuba, Paraná290[23] as Brachycephalus sp. 1; [27] as Brachycephalus sp. nov. 1; [72] as B. sulfuratus; [115] as Brachycephalus aff. hermogenesi]; [116] as B. hermogenesi
Brachycephalus sp. (cf. B. sulfuratus)didactylusDona Francisca (26°09’52” S, 48°59’23” W), municipality of Joinville, Santa Catarina150[23] as Brachycephalus sp. 1; [27] as Brachycephalus sp. nov. 1
Brachycephalus sp. (B. sulfuratus or B. hermogenesi)didactylusEstação Ecológica Juréia-Itatins (c. 24°27’ S, 47°24’ W), municipality of Iguape, São Paulo?[63] as B. hermogenesi
Brachycephalus sp. (cf. B. sulfuratus)didactylusEstrada do Rio do Júlio (26°17’02” S, 49°06’08” W), municipality of Joinville, Santa Catarina650[23] as Brachycephalus sp. 1; [117] as Brachycephalus sp.
Brachycephalus sp. (cf. B. sulfuratus)didactylusFazenda Pico Paraná (25°13’29” S, 48°51’17” W), municipality of Campina Grande do Sul, Paraná1050–1085[23] as Brachycephalus sp. 1; [27] as Brachycephalus sp. nov. 1
Brachycephalus sp. (cf. B. sulfuratus)didactylusFazenda Primavera (24°53’08” S, 48°45’51” W), municipality of Tunas do Paraná, Paraná1060[23] as Brachycephalus sp. 1; [27] as Brachycephalus sp. nov. 1
Brachycephalus sp. (B. sulfuratus or B. hermogenesi)didactylusMunicipality of Ibiúna (c. 23°39’ S, 47°13’ W), São Paulo?[72] as B. hermogenesi
Brachycephalus sp. (B. sulfuratus or B. hermogenesi)didactylusMunicipality of Juquitiba (c. 23°56’ S, 47°04’ W), São Paulo?[63] as B. hermogenesi; [72] as B. hermogenesi
Brachycephalus sp. (cf. B. hermogenesi)didactylusMunicipality of Paraty (c. 23°13’07” S, 44°43’15” W), Rio de Janeiro?[18] as B. hermogenesi
Brachycephalus sp. (B. sulfuratus or B. hermogenesi)didactylusMunicipality of Peruíbe (24°18’ S, 46°59’ W), São Paulo?[72] as B. hermogenesi
Brachycephalus sp. (B. sulfuratus or B. hermogenesi)didactylusMunicipality of Piedade (c. 23°54’S, 47°25’ W), São Paulo?[81] as B. hermogenesi; [118] as B. hermogenesi
Brachycephalus sp. (B. sulfuratus or B. hermogenesi)didactylusMunicipality of Registro (c. 24°30’ S, 47°51’ W), São Paulo?[72] as B. hermogenesi
Brachycephalus sp. (B. sulfuratus or B. hermogenesi)didactylusMunicipality of Ribeirão Grande (c. 24°06’ S, 48°22’ W), São Paulo?[63] as B. hermogenesi
Brachycephalus sp. (B. sulfuratus or B. hermogenesi)didactylusMunicipality of Tapiraí (c. 23°57’55” S, 47°30’19” W), São Paulo870[63] as B. hermogenesi; [118] as B. hermogenesi
Brachycephalus sp. (B. sulfuratus or B. hermogenesi)didactylusParque Estadual de Jacupiranga (c. 24°38’ S, 48°24’ W), municipality of Eldorado, São Paulo?[72] as B. hermogenesi
Brachycephalus sp. (B. hermogenesi or B. sulfuratus)didactylusParque Natural Municipal Nascentes de Paranapiacaba (23°46’10” S, 46°17’36” W), municipality of Santo André, São Paulo840[119] as B. hermogenesi
Brachycephalus sp. (cf. B. sulfuratus)didactylusPico Agudinho (25°36’24” S, 48°43’33” W), Serra da Prata, municipality of Morretes, Paraná385[23] as Brachycephalus sp. 1; [27] as Brachycephalus sp. nov. 1
Brachycephalus sp. (B. sulfuratus or B. hermogenesi)didactylusReserva Betary (24°33’08” S, 48°40’49” W), municipality of Iporanga, São Paulo190This study
Brachycephalus sp. (B. hermogenesi or B. sulfuratus)didactylusReserva Biológica do Alto da Serra de Paranapiacaba (23°46’40” S, 46°18’45” W), municipality of Santo André, São Paulo800–850[23] as B. hermogenesi; [63] as B. hermogenesi; [119] as B. hermogenesi
Brachycephalus sp. (B. sulfuratus or B. hermogenesi)didactylusReserva Florestal de Morro Grande (23°42’08” S, 46°58’22” W), municipality of Cotia, São Paulo990?[23] as B. hermogenesi, [63] as B. hermogenesi; [72] as B. hermogenesi; [94] as B. hermogenesi
Brachycephalus sp. (cf. B. sulfuratus)didactylusSítio Ananias (25°47’08” S, 48°43’03” W), municipality of Guaratuba, Paraná25[23] as Brachycephalus sp. 1; [27] as Brachycephalus sp. nov. 1
Brachycephalus sp.ephippiumParanapiacaba (23°46’30” S, 46°17’57” W), municipality of Santo André, São Paulo825[27] as Brachycephalus sp. 1; [66] as B. ephippium
Brachycephalus sp.ephippiumParque Natural Municipal Nascentes de Paranapiacaba (23°46’10” S, 46°17’36” W), municipality of Santo André, São Paulo800–1164?[120] as Brachycephalus sp.
Brachycephalus sp.ephippiumPenísula do Bororé (23°47’11” S, 46°38’45” W), Represa Billings, Grajaú District, municipality of São Paulo, São Paulo780[27] as Brachycephalus nodoterga; [99] as another species than B. nodoterga of [27]
Brachycephalus sp.ephippiumReserva Biológica do Alto da Serra de Paranapiacaba (23°46’40” S, 46°18’45” W), municipality of Santo André, São Paulo800[27] as Brachycephalus sp. 1
Brachycephalus sp.ephippiumTheodoro de Oliveira (first position: 22°22’11” S, 42°33’25” W), Parque Estadual dos Três Picos, municipality of Nova Friburgo, Rio de Janeiro1100–1200[67] as Brachycephalus sp.; [95] as Brachycephalus sp. nov.
Brachycephalus sp.pernixPedra Branca do Araraquara (25°56’00” S, 48°52’50” W), Serra do Araraquara, municipality of Guaratuba, Paraná1000[23] as Brachycephalus sp. 5
Brachycephalus sp.pernixPico Paraná (25°15′10″ S, 48°48′32″ W), Serra dos Órgãos, municipality of Antonina, Paraná1880This study
Brachycephalus sp.pernixSerra Canasvieiras (25°36’58” S, 48°46’59” W), boundary of the municipalities of Guaratuba and Morretes, Paraná1080[23] as Brachycephalus sp. 5; [25] as B. sp. Canasvieiras; [28] as not identified; [73] without species identification
Brachycephalus sp.pernixTupipiá (25°15’13” S, 48°48’20” W), Serra dos Órgãos, municipality of Antonina, Paraná1560This study, [27] as B. brunneus; [28] as B. brunneus; [29] as B. sp. Tupipiá, [73] without species identification
Brachycephalus sp. (cf. B. darkside juvenile)?Parque Estadual da Serra do Brigadeiro (cf. 20°43′16″ S, 42°29′05″ W), municipality of Araponga, Minas Gerais1330?[85] as Brachycephalus cf. didactylus
1 Data with “?” were not available in literature.
Table 2. Status of conservation of identified species of Brachycephalus according to IUCN [48]. Abbreviations: EO = extent of occurrence (see text for details); MMA = Ministério do Meio Ambiente (Brazil).
Table 2. Status of conservation of identified species of Brachycephalus according to IUCN [48]. Abbreviations: EO = extent of occurrence (see text for details); MMA = Ministério do Meio Ambiente (Brazil).
Species 1Localities 1Altitudinal Range (m a.s.l.) 1,2Evaluation of EO (ha)Flow Chart Pathway 3PopulationConservation Status—Criteria
Previous 2This Study 2Locations 2Individuals 2, 3IUCNMMAOthersThis Study
B. didactylus group
B. didactylus835–1110---702,983.45479,655,049LC [41]------VU - B1ab(i,iii)
B. hermogenesi110–1090567,589.9 [23]143,325.051?LC [33]------VU - B1ab(i,iii)
B. pulex1800–930488.2 [23]482.311?---------VU - D2
B. sulfuratus2640–1205778,458.4 [23]3,021,786.151302,178,610---------LC
B. ephippium group
B. alipioi31070–110038,950.0 [47], 27,930.0 [43]1,706.131?DD [35]NT [43]---CR - B1ab(i,iii)
B. bufonoides1?---?4??---------DD
B. crispus1800–1190??21?---------DD
B. darkside21265–1500--5,700.811?---------CR - B1ab(i,iii)
B. ephippium31200–1250?1,792,535.15613,336,461LC [38]------VU - B1ab(i,iii)
B. garbeanus71130–190012,268.0 [23]6,426.552?---------EN - B1ab(i,iii)+2ab(ii,iii)
B. guarani1500–900??21?---------DD
B. margaritatus6600–98018,272.9 [23]10,710.552?---------EN - B1ab(i,iii)
B. nodoterga5700–9009,690.0 [47], 108,280.0 [42]28.458.153?DD [31]DD [42]---VU - B1ab(i,iii)
B. pitanga4900–11402,377.1 [23]2,245.15129,157,136---------LC
B. toby1750–1060??21?---------DD
B. vertebralis2760–1110161,990.0 [47], 18,580.0 [44]?22?DD [37]DD [44]---DD
B. pernix group
B. actaeus720–530---15,841.662?---------EN - B1ab(i,iii)+2ab(ii,iii)
B. albolineatus4500–83534.4 [74]2,784.45121,076,087------DD [74]VU - B1ab(i,iii)
B. auroguttatus11070–1100---?31 ---------DD
B. boticario1685–79511.1 [23]38.811?---------CR - B1ab(i,iii)+2ab(ii,iii)
B. brunneus61095–17701,100.0 [47], 5,687.1 [23]5,385.662?DD [39]------LC
B. coloratus11145–1230---37.411 ------DD [50]VU - D2
B. curupira31095–13202,211.54 [23]4,751.46221,117,312------DD [50]LC
B. ferruginus1965–1,47038,950.0 [47], 5,475.5 [23]5,994.311?DD [34]------LC
B. fuscolineatus2525–79023.63 [23], 23.8 [24]23.812119,000---------CR - B1ab(i,iii)+2ab(ii,iii)
B. izecksohni1980–13401,100.0 [47], 350.4 [23]378.311?DD [40]------VU - D2
B. leopardus21340–1645176.7 [23]363.113?---------EN - B1ab(i,iii)+2ab(ii,iii)
B. mariaeterezae11265–1270---?31?---------DD
B. mirissimus1470–54056.8 [25]56.81178,344------CR [25]CR - B1ab(i,iii)+2ab(ii,iii)
B. olivaceus4650–98512,531.6 [23]18,850.152?---------EN - B1ab(i,iii)+2ab(ii,iii)
B. pernix11135–14051,950.0 [47], 432.1 [23], 400 [45]389.411?DD [32]CR - B1ab(iii)+2ab(iii) [45]---VU - D2
B. pombali2845–130031,300.0 [47]?21?DD [36]------DD
B.quiririensis21240–13801339.0 [23]629.011?---------CR - B1ab(i,iii)+2ab(ii,iii)
B. tridactylus1805–91041.4 [23]41.411?---------VU - D2
B. verrucosus1455–945---?21?---------DD
Incertae sedis
B. atelopoide1?---?4??---------DD
1 According Table 1.2 Data with “?” could not be estimate because of lack of information in literature. 3 According Figure 1.

Share and Cite

MDPI and ACS Style

Bornschein, M.R.; Pie, M.R.; Teixeira, L. Conservation Status of Brachycephalus Toadlets (Anura: Brachycephalidae) from the Brazilian Atlantic Rainforest. Diversity 2019, 11, 150. https://doi.org/10.3390/d11090150

AMA Style

Bornschein MR, Pie MR, Teixeira L. Conservation Status of Brachycephalus Toadlets (Anura: Brachycephalidae) from the Brazilian Atlantic Rainforest. Diversity. 2019; 11(9):150. https://doi.org/10.3390/d11090150

Chicago/Turabian Style

Bornschein, Marcos R., Marcio R. Pie, and Larissa Teixeira. 2019. "Conservation Status of Brachycephalus Toadlets (Anura: Brachycephalidae) from the Brazilian Atlantic Rainforest" Diversity 11, no. 9: 150. https://doi.org/10.3390/d11090150

Note that from the first issue of 2016, MDPI journals use article numbers instead of page numbers. See further details here.

Article Metrics

Back to TopTop