Description and evolution of the larynx of the Physalaemus olfersii species group, with remarks on the laryngeal anatomy of the P. cuvieri clade (Amphibia: Anura: Leiuperinae).

The anuran larynx is an organ of great evolutionary interest because it impacts male reproductive success in courtships. However, little is known about the diversity of the larynx's anatomy, evolutionary history and systematics importance. Here, we describe and compare the anatomy of the larynx of 10 Physalaemus species of the P. cuvieri clade, focusing on the P. olfersii species group. We also reconstructed the ancestral states and tested the phylogenetic signal for the anatomical features. In all the species, the larynx has a general globular shape with the arytenoid cartilages covering almost its entire dorsal surface, while the anterior process of the cricoid cartilages covers most of the ventral surface. The size of the secondary fibrous mass, the thickness of the vocal membrane, and the attachment position of the vocal membrane's free edge considerably differ among the species. Moreover, only four species of a single clade in the P. olfersii species group have the primary fibrous mass well-developed with a suspended region in the dorsolateral passage. We found a significant phylogenetic signal for all these characters. Ancestral reconstructions pointed to reduction tendencies in the thickness of the vocal membrane and the size of the secondary fibrous mass, and a shift of the ventral attachment of the vocal membrane, increasing the angle of its free edge along the phylogeny. This latter trait can diagnose the entire Physalaemus olfersii group, which has the ventral ends of the arytenoids positioned posteriorly, giving this group the steepest angles for the vocal membrane's free edge in relation to the frontal plane. Based on our results, the larynges can contribute to the Physalaemus olfersii species group's systematics and could be elucidative to understand the evolution of the genus. High levels of anatomical and bioacoustical complexity and diversity observed in the group support the expected correlation between vocal anatomy and bioacoustical signal.

How long does it take to know a species? Redescription, vocalization, and chigger mites infestation pattern in the centenary emBrachycephalus/em emgarbeanus/em (Anura: Brachycephalidae).

Although we celebrate the centennial of Brachycephalus garbeanus' discovery, little progress has been done on understanding this species' biology apart from a few morphological and ecological studies, which includes its redescription based on three specimens from the type-series, microhabitat use, sexual dimorphism in body size, and feeding habits. This species is endemic to the Serra dos Órgãos Mountain range, in the state of Rio de Janeiro, southeastern Brazil. Here we redescribe B. garbeanus based on a wide sampling, including its advertisement and aggressive calls, and also the chigger mites infestation pattern. The advertisement call is longer than 25.8 s with pulsed notes series emitted at an average rate of 2.3 notes/s and 14.1 pulses/s; long inter-note interval with 320 ms; notes with distinctly short pulses (1 to 16 ms); low dominant frequency for this genus (3.0-5.4 kHz) and presence of four harmonics. This species is often parasitized by chigger mites of Hannemania, with a prevalence of infection of 67%, mainly affecting the ventral body surface. Females had a higher prevalence of parasites than males and there was no correlation found between the size of specimens and the number of parasites. Our study, provides an important and overdue taxonomical contribution, including a large amount of novel information for B. garbeanus.

A new species of the Scinax catharinae species group (Anura: Hylidae) from the Atlantic rainforest of northeastern Minas Gerais, southeastern Brazil.

We describe a new species of the Scinax catharinae group from one of the last remaining fragments of Atlantic Forest in northeastern Minas Gerais. The new species is distinguished from most species of the group mainly by having the mucronate snout in dorsal view, the unique light-yellow coloration on the gular surface in life contrasting with its overall color pattern, and the m. depressor mandibulae with an origin on the dorsal fascia at the level of the m. dorsalis scapulae. These two latter features are novelties in the diagnosis of Scinax species. The new species is currently known only from municipality of Santa Maria do Salto, state of Minas Gerais, Brazil. This locality is a remaining fragment of Atlantic Forest and has a peculiar geomorphic feature that has been poorly sampled so far.

Rediscovery of the toadlet Brachycephalus bufonoides Miranda-Ribeiro, 1920 (Anura: Brachycephalidae) with osteological and acoustic descriptions.

Brachycephalus bufonoides was described as a "variety" of B. ephippium based on two specimens which 90 years later was considered full species. Besides its brief original description, nothing else is known for this species. Herein we report the rediscovery of the pumpkin-toadlet Brachycephalus bufonoides from Nova Friburgo, State of Rio de Janeiro, the second most populous area within the Atlantic Forest in Brazil. A detailed osteological description of this species was also provided, including skull, hyolaryngeal skeleton and postcranium skeleton. The laryngeal skeleton of Brachycephalus genus was depicted for the first time. We conducted a molecular phylogenetic analysis of Brachycephalus using DNA sequences comprising two fragments of mitochondrial gene (16S). Both analysis with Bayesian inference and maximum parsimony supported the recognition of B. bufonoides as an exclusive lineage, allocated within the B. ephippium species group in B. vertebralis lineage. We improved the diagnosis and variation of the species, including more collected specimens, coloration in vivo and advertisement call description. Compared with its congeners, B. bufonoides has skin on head and dorsum with dermal hyperossification; skull with hyperossification of postorbital crests; a pair of hyperossified bulges about equidistant between postorbital crests; fourth presacral vertebra with transverse process hyperossified, ornamented and sacral diapophyses hyperossified, which can be seen externally (lineage of B. vertebralis sensu Condez et al. 2020); presence of dermal ossification as separated bulges of each vertebrae; general background color orange with different intensities of dark orange blotches on dorsum, including bordering of sacral region; absence of osteoderms and presence of warts on the dorsolateral surface of body; medium body size (SVL of adults: 12.0-14.5 mm for males and 14.7-16.3 mm for females; Table 1); rough dorsum; advertisement calls with 13 to 17 pulses; presence of pulse period modulation; and advertisement calls with notes longer than 0.2 s (0.22 to 0.31 s). Herein an important contribution for the taxonomy and systematics of this genus is provided, including a large amount of novel information for B. bufonoides from different sources (i.e., molecular, morphological variation, bioacoustic), allowing it to be included in future studies of species delimitation and relationships within Brachycephalus. Also, the discovery of this species reiterates the importance of Nova Friburgo for the conservation of the Atlantic Forest biodiversity.

Review of bioacoustical traits in the genus Physalaemus Fitzinger, 1826 (Anura: Leptodactylidae: Leiuperinae).

Given the importance of acoustic communication in intraspecific recognition during mating activity, acoustic traits have been widely used to clarify the taxonomy of anurans. They have been particularly useful in the study of taxa with high morphological similarity such as the Neotropical genus Physalaemus. Here, we reviewed the acoustic repertoires of the species of Physalaemus based on homology hypotheses in order to make comparisons more properly applicable for taxonomic purposes. We covered all the known clades and species groups for the genus, analyzing 45 species (94 % of the currently recognized taxa). Different call types were labeled with letters (i.e., A, B, and C) to avoid speculative functional propositions for the call types. In order to identify correctly the observed frequency bands, we propose a method to interpret them based on the predicted graphic behavior on audiospectrogram and on the mathematic relationship among bands considering each kind of band production (e.g., harmonics and sidebands). We found different acoustic traits between the major clades P. signifer and P. cuvieri. Species in the P. signifer clade have more than one call type (67 % of species in the clade). Furthermore, all species of this clade have A calls with pulses and/or low fundamental frequency (< 500 Hz). In the P. cuvieri clade, species emit only one call type and, in most species, this call is a continuous whine-like emission with relatively high fundamental frequency (> 400 Hz) and several S-shaped harmonics (except for species of P. henselii and P. olfersii groups, P. centralis, and P. cicada). Within the P. signifer clade, pulsed calls are present in P. angrensis, P. atlanticus, P. bokermanni, P. crombiei, P. irroratus, P. moreirae, P. nanus, and P. obtectus, whereas within the P. cuvieri clade this feature is restricted to a few species (10 % of the clade): P. jordanensis, P. feioi, and P. orophilus. A principal component analysis of the quantitative data indicates two clusters that substantially correspond to the composition of these two major clades with a few exceptions. Overall, the cluster composed of taxa of the P. signifer clade has lower fundamental frequency, bandwidth and dominant frequency at the end of the call and higher frequency delta and dominant frequency at the end of the call than the cluster with most taxa of the P. cuvieri clade. We also identified and described several similarities among acoustic signals of closely related species, which might correspond to synapomorphies in the evolution of the acoustic signal in the group. Species of the P. deimaticus group emit long sequences of very short A calls with low fundamental frequency (< 300 Hz) and short duration (< 0.2 s). Most species in the P. signifer group have clearly pulsed calls and emit at least two different call types. Species in the P. henselii group have calls with only high frequency bands (> 1700 Hz). Species in P. cuvieri group have continuous calls that resemble nasal-like sounds or whines, with downward frequency modulation. Species in the P. olfersii group emit long calls (> 1 s) with ascendant and periodic frequency modulation. Calls of the species in the P. biligonigerus and P. gracilis groups usually have continuous whine-like calls with call envelopes very variable within species. In addition, we describe traits in the genus for the first time, such as complex traits not predicted by simple and linear acoustic models (nonlinear phenomena), and discuss the application of acoustic traits to taxonomy and phylogenetics and morphological constraints of the vocal apparatus that might be related to the different acoustic properties found.

A new species of the Ololygon catharinae species group (Anura: Hylidae) from the Cerrado biome, State of Goiás, Central Brazil.

We describe a new species of Ololygon from the Cerrado biome, Central Brazil. The new species is assigned to the Ololygon catharinae species group based in morphological attributes. Ololygon goya sp. nov. in known only from type locality-Sítio d'Abadia municipality, State of Goiás-and associated with riparian environments connected to seasonal dry forest on the banks of the Corrente river, a tributary of the Paranã river basin. The new species is characterized by the combination of the following characters: medium size (24.4-38.8 mm SVL); snout subovoid in dorsal view; canthus rostralis well defined; males with hypertrophied forearms and nuptial pads; inguinal region and hidden surfaces of thigh with irregular dark brown spots on pale yellowish background. We also describe the external morphology and oral morphology of tadpoles, and the advertisement calls of the new species.

Timeless standards for species delimitation.

Recently a new species of bombyliid fly, Marleyimyia xylocopae, was described by Marshall & Evenhuis (2015) based on two photographs taken during fieldwork in the Republic of South Africa. This species has no preserved holotype. The paper generated some buzz, especially among dipterists, because in most cases photographs taken in the field provide insufficient information for properly diagnosing and documenting species of Diptera.

The First Bromeligenous Species of Dendropsophus (Anura: Hylidae) from Brazil's Atlantic Forest.

We describe a new treefrog species of Dendropsophus collected on rocky outcrops in the Brazilian Atlantic Forest. Ecologically, the new species can be distinguished from all known congeners by having a larval phase associated with rainwater accumulated in bromeliad phytotelms instead of temporary or lentic water bodies. Phylogenetic analysis based on molecular data confirms that the new species is a member of Dendropsophus; our analysis does not assign it to any recognized species group in the genus. Morphologically, based on comparison with the 96 known congeners, the new species is diagnosed by its small size, framed dorsal color pattern, and short webbing between toes IV-V. The advertisement call is composed of a moderate-pitched two-note call (~5 kHz). The territorial call contains more notes and pulses than the advertisement call. Field observations suggest that this new bromeligenous species uses a variety of bromeliad species to breed in, and may be both territorial and exhibit male parental care.

Phylogenetic systematics of egg-brooding frogs (Anura: Hemiphractidae) and the evolution of direct development.

Egg-brooding frogs (Hemiphractidae) are a group of 105 currently recognized Neotropical species, with a remarkable diversity of developmental modes, from direct development to free-living and exotrophic tadpoles. Females carry their eggs on the back and embryos have unique bell-shaped gills. We inferred the evolutionary relationships of these frogs and used the resulting phylogeny to review their taxonomy and test hypotheses on the evolution of developmental modes and bell-shaped gills. Our inferences relied on a total evidence parsimony analysis of DNA sequences of up to 20 mitochondrial and nuclear genes (analyzed under tree-alignment), and 51 phenotypic characters sampled for 83% of currently valid hemiphractid species. Our analyses rendered a well-resolved phylogeny, with both Hemiphractidae (sister of Athesphatanura) and its six recognized genera being monophyletic. We also inferred novel intergeneric relationships [((Cryptobatrachus, Flectonotus), (Stefania, (Fritziana, (Hemiphractus, Gastrotheca))))], the non-monophyly of all species groups previously proposed within Gastrotheca and Stefania, and the existence of several putative new species within Fritziana and Hemiphractus. Contrary to previous hypotheses, our results support the most recent common ancestor of hemiphractids as a direct-developer. Free-living aquatic tadpoles apparently evolved from direct-developing ancestors three to eight times. Embryos of the sister taxa Cryptobatrachus and Flectonotus share a pair of single gills derived from branchial arch I, while embryos of the clade including the other four genera have two pairs of gills derived from branchial arches I and II respectively. Furthermore, in Gastrotheca the fusion of the two pairs of gills is a putative synapomorphy. We propose a revised taxonomy concordant with our optimal topologies.

A new species of the Scinax catharinae Group (Anura: Hylidae) from Northeastern Brazil.

We describe a new species of the Scinax catharinae Group from Municipality of Porto Seguro, State of Bahia northeastern Brazil. The new species is mainly characterized by its small size, nuptial pad dark colored, and compound pectoral fold. Additionally, we describe the structure of its nuptial pad and compare it with that of S. agilis. We also briefly discuss its phylogenetic relationships within Scinax. 

High levels of diversity uncovered in a widespread nominal taxon: continental phylogeography of the neotropical tree frog Dendropsophus minutus.

Species distributed across vast continental areas and across major biomes provide unique model systems for studies of biotic diversification, yet also constitute daunting financial, logistic and political challenges for data collection across such regions. The tree frog Dendropsophus minutus (Anura: Hylidae) is a nominal species, continentally distributed in South America, that may represent a complex of multiple species, each with a more limited distribution. To understand the spatial pattern of molecular diversity throughout the range of this species complex, we obtained DNA sequence data from two mitochondrial genes, cytochrome oxidase I (COI) and the 16S rhibosomal gene (16S) for 407 samples of D. minutus and closely related species distributed across eleven countries, effectively comprising the entire range of the group. We performed phylogenetic and spatially explicit phylogeographic analyses to assess the genetic structure of lineages and infer ancestral areas. We found 43 statistically supported, deep mitochondrial lineages, several of which may represent currently unrecognized distinct species. One major clade, containing 25 divergent lineages, includes samples from the type locality of D. minutus. We defined that clade as the D. minutus complex. The remaining lineages together with the D. minutus complex constitute the D. minutus species group. Historical analyses support an Amazonian origin for the D. minutus species group with a subsequent dispersal to eastern Brazil where the D. minutus complex originated. According to our dataset, a total of eight mtDNA lineages have ranges >100,000 km2. One of them occupies an area of almost one million km2 encompassing multiple biomes. Our results, at a spatial scale and resolution unprecedented for a Neotropical vertebrate, confirm that widespread amphibian species occur in lowland South America, yet at the same time a large proportion of cryptic diversity still remains to be discovered.

The identity of Rana margaritifera Laurenti, 1768 (Anura, Bufonidae).

Rana margaritifera was described by Laurenti in 1768 and currently is associated to the genus Rhinella, under the combination Rhinella margaritifera. Currently, the R. margaritifera species group consists of 16 recognized species. Furthermore, many additional species have been suggested to exist in this group which highlights the ambiguity surrounding the identity of Rhinella margaritifera and impend further description of the species in this group. After an exhaustive bibliographic review, we concluded that the recent designation of a lectotype for R. margaritifera is invalid according with Art. 73, ICZN, 1999. Herein, we designate and provide the description of a neotype for Rana margaritifera Laurenti, 1768.

Nomenclatural issues in ornithology: the incredible controversy on the identity of a long overlooked Brazilian bird.

The identity of Scytalopus speluncae (Ménétriés, 1835) (Aves: Passeriformes: Rhinocryptidae), a tapaculo from southeastern Brazil, has been the matter of debate during the last eight years. A group of ornithologists considers that the nomen Scytalopus speluncae should be attributed to a species endemic to coastal mountains of southeastern Brazil, whereas another group considers it a species from the drier environments of another mountain belt in Minas Gerais, southeastern Brazil. Both research groups disagree on the identity of the still extant but damaged alleged holotype, deposited at the Zoological Institute of the Russian Academy of Sciences, St. Petersburg, on the identity of the holotype specimen illustration from a plate accompanying the species description, and even on the type locality. To further complicate this matter of identity, members of each research group, based on their own interpretations of the identity of Scytalopus speluncae, described and named again the two species with different nomina, erecting at least one unnecessary nomen. After almost ten years of a debate, there is still no consensus on the identity of the species, and there are now at least three available nomina for apparently only two distinct biological species. As taxonomists belonging to fields of zoology other than ornithology, and realizing the above situation is mainly a nomenclatural one, we herein present a summary of the contentious issue, try to distinguish what seems to be facts and speculation and based on these we consider the rules of the International Code of Zoological Nomenclature (the Code) whenever appropriate, in the hope of bringing some objectivity to the debate. We conclude that no unequivocal evidence was presented to decide to which species the type specimen belongs solely based on its morphological characters, since the holotype presents considerable damage. On the other hand, the original designation of the type locality by Ménétriés (1835) as São João del Rei, in southeastern Brazil, is unquestionable. Thus, the posterior re-designation of the type locality by a group of ornithologists was not done according to the Code. Contrary to the proposition made by a group of ornithologists that the nomen Scytalopus speluncae should be maintained to the species so called up to 2006 until further evidence confirms the identity of the holotype, we here argue that for now the nomen should be applied to the only species known to occur in the vicinities of the type locality, as established by the author of the species. Nevertheless, we also agree that obtaining molecular data from the holotype should be attempted to definitely solve the query.