A new species of the Scinax cruentomma group (Anura: Hylidae) from the Ucayali River basin of Loreto, Peru.

We describe a new species of the Scinax cruentomma species group, with a red streak in the iris and a weakly bilobate vocal sac. It is known from oligotrophic soils in the sedimentary basin of the Ucayali River near Jenaro Herrera (province of Requena, Peru) and Ro Blanco (buffer zone of the Matses Indigenous territory and reserve). The new species can be distinguished from the other species of the S. cruentomma group by its small snout-vent length, body and iris color patterns, weakly bilobate vocal sac, myological characters, and the number of notes and pulses of the advertisement call. It is morphologically most similar to S. strussmannae, from which the advertisement call, nostril, canthus rostralis, and loreal region can distinguish it.

Lost in time: Unraveling the identity of Vitreorana parvula (Boulenger, 1895) (Anura: Centrolenidae).

Vitreorana parvula was the first glassfrog described for the Atlantic Forest. The species, however, has become a taxonomic puzzle as the only known individual is the lectotype from the 19th century, which is not particularly well-preserved or accompanied by a detailed original description. To solve this problem, we collected topotypic specimens, as well as advertisement calls, tissue samples, and natural history data, and compared them to other Vitreorana species. Our results show clear morphological, acoustic, and genetic differences between V. parvula and other species of Vitreorana, except for V. uranoscopa. Following our results, we consider V. uranoscopa as a junior synonym of V. parvula and redescribe the species based on topotypic material, while summarizing relevant variation from across its distribution.

A new dwarf gecko of the genus Pseudogonatodes (Squamata: Sphaerodactylidae) from the eastern slope of the Andean Cordillera de Mrida in northern South America.

Pseudogonatodes is a poorly known genus of small bodied, diurnal, ground-dwelling geckos widely distributed in northern South America. No additional species have been described in over two decades. Herein we describe a new species from the eastern slope of the Cordillera de Mrida in the Venezuelan Andes. The new species is readily diagnosable morphologically from the other seven recognized species of Pseudogonatodes by having a single postnasal scalea putative autapomorphy. Furthermore, it is characterized by a unique combination of phenotypic characters that includes granular dorsal scales, three large postrostrals, five to six loreals, mental U-shaped, four to six postmentals, 2629 ventrals between anterior levels of fore- and hind limbs, third lamella under fourth toe not distinctly enlarged, and subcaudal pattern 11. In addition to characterizing the external morphology, we present a description of the skull, based on 3D digital models reconstructed from high resolution computed microtomography scans. The discovery of this new species highlights the still underestimated diversity of this group of Neotropical dwarf geckos and underscores the need for further studies on its systematics and taxonomy.

Multiple Routes to Color Convergence in a Radiation of Neotropical Poison Frogs.

Convergent evolution is defined as the independent evolution of similar phenotypes in different lineages. Its existence underscores the importance of external selection pressures in evolutionary history, revealing how functionally similar adaptations can evolve in response to persistent ecological challenges through a diversity of evolutionary routes. However, many examples of convergence, particularly among closely related species, involve parallel changes in the same genes or developmental pathways, raising the possibility that homology at deeper mechanistic levels is an important facilitator of phenotypic convergence. Using the genus Ranitomeya, a young, color-diverse radiation of Neotropical poison frogs, we set out to (1) provide a phylogenetic framework for this group, (2) leverage this framework to determine if color phenotypes are convergent, and (3) to characterize the underlying coloration mechanisms to test whether color convergence occurred through the same or different physical mechanisms. We generated a phylogeny for Ranitomeya using ultraconserved elements and investigated the physical mechanisms underlying bright coloration, focusing on skin pigments. Using phylogenetic comparative methods, we identified several instances of color convergence, involving several gains and losses of carotenoid and pterin pigments. We also found a compelling example of nonparallel convergence, where, in one lineage, red coloration evolved through the red pterin pigment drosopterin, and in another lineage through red ketocarotenoids. Additionally, in another lineage, "reddish" coloration evolved predominantly through structural color mechanisms. Our study demonstrates that, even within a radiation of closely related species, convergent evolution can occur through both parallel and nonparallel mechanisms, challenging the assumption that similar phenotypes among close relatives evolve through the same mechanisms.

An elusive giant: A new species of Vitreorana Guayasamin et al., 2009 (Anura: Centrolenidae) from the northern Atlantic Forest with an osteological description and comments on integumentary spicules.

We describe a new species of Vitreorana from the Atlantic Forest of southern Bahia state, in north-eastern Brazil. Vitreorana assuh sp. nov. is by far the largest species of the genus (snout-to-vent length [SVL] = 30.9 mm and 34.1 mm in one male and female, respectively; maximum SVL recorded for all other species = 28.0 mm). The new species is morphologically most similar to V. franciscana, from the Cerrado biome in Brazil. Besides its large size, the new species has, among other diagnostic characters, the presence of vomerine teeth and the occurrence of enameled iridophores in the pericardium, hepatic and urinary bladder peritonea. Furthermore, we describe and compare the complete mineralized skeleton of the new species and V. uranoscopa, and describe and discuss the variation of integumentary spicules in all species of the genus and their relevance to Vitreorana systematics.

A new species of Chiasmocleis Méhely, 1904 (Anura: Microhylidae) from the Peruvian Amazonia.

We describe a new species of Chiasmocleis from the Amazonian forest of Peru. The new species is characterized by its medium size (snout-to-vent length = 18.2-20.8 mm in females, and 16.5 mm in one male), hands and feet with slightly developed fridges in females (more developed in male), and presence of a femoral line in all individuals. We also infer its phylogenetic position using DNA sequences of fragments of the mitochondrial genes for 16S rRNA and cytochrome oxidase subunit 1 (COI), and describe the mineralized skeleton through 3D models generated by computed tomography (CT-scan). Based on our results, we discuss the variation of some osteological characters traditionally used in the systematics of the genus.

A new species of nurse-frog (Aromobatidae, Allobates) from the Amazonian forest of Loreto, Peru.

We describe a new species of nurse-frog (Aromobatidae, Allobates) from the Amazonian forest of Loreto, Peru using morphological, acoustic and genetic data. Our phylogenetic analysis placed Allobates sieggreenae sp. nov. as the sister species of A. trilineatus, the most similar-looking species and with which it was previously confused. However, the new species has a brown dorsum, solid dark brown lateral dark stripe not fading towards groin, adult males with few and sparse melanophores over a cream background on chin, chest, and belly, dark transverse bars absent on thighs, and an advertisement call formed by a trill of single notes (in A. trilineatus dorsum dark brown, blackish brown lateral dark stripe, paler from mid-body to groin, adult males with a dark background color on chin, chest, and belly due to a dense layer of melanophores, dark transverse bar present on dorsal surface of thighs, and trills of paired notes). Allobates sieggreenae is known from two localities of Amazonian white-sand forest ecosystems east of the Ucayali River.

Total evidence and sensitivity phylogenetic analyses of egg-brooding frogs (Anura: Hemiphractidae).

We study the phylogenetic relationships of egg-brooding frogs, a group of 118 neotropical species, unique among anurans by having embryos with large bell-shaped gills and females carrying their eggs on the dorsum, exposed or inside a pouch. We assembled a total evidence dataset of published and newly generated data containing 51 phenotypic characters and DNA sequences of 20 loci for 143 hemiphractids and 127 outgroup terminals. We performed six analytical strategies combining different optimality criteria (parsimony and maximum likelihood), alignment methods (tree- and similarity-alignment), and three different indel coding schemes (fifth character state, unknown nucleotide, and presence/absence characters matrix). Furthermore, we analyzed a subset of the total evidence dataset to evaluate the impact of phenotypic characters on hemiphractid phylogenetic relationships. Our main results include: (i) monophyly of Hemiphractidae and its six genera for all our analyses, novel relationships among hemiphractid genera, and non-monophyly of Hemiphractinae according to our preferred phylogenetic hypothesis; (ii) non-monophyly of current supraspecific taxonomies of Gastrotheca, an updated taxonomy is provided; (iii) previous differences among studies were mainly caused by differences in analytical factors, not by differences in character/taxon sampling; (iv) optimality criteria, alignment method, and indel coding caused differences among optimal topologies, in that order of degree; (v) in most cases, parsimony analyses are more sensitive to the addition of phenotypic data than maximum likelihood analyses; (vi) adding phenotypic data resulted in an increase of shared clades for most analyses.

Integrative taxonomy reveals a new but common Neotropical treefrog, hidden under the name Boana xerophylla.

Boana xerophylla is a common treefrog widely distributed in northern Brazil, Colombia, French Guiana, Guyana, Panama, Suriname, Trinidad and Tobago, and Venezuela. A recent study found molecular, acoustic, and morphometric differences between the populations located on opposite sides of the Orinoco River. Here, we carry out an updated molecular phylogenetic analysis, including new samples from all the countries along the distribution area, and analyzed additional call recordings from Brazil, Colombia, French Guiana, Panama, Trinidad and Tobago, and Venezuela. Our phylogenetic inference reveals three geographically restricted lineages: one in the eastern Guiana Shield (corresponding to B. xerophylla sensu stricto), another in the western Guiana Shield, and a third one north of the Orinoco River. Morphological and acoustic data agree with the differentiation between the populations north of the Orinoco River and the eastern Guiana Shield despite the low genetic p-distances observed (16S rRNA: 0.72.2 %). We argue that the populations north of the Orinoco River correspond to a new species, sister of B. xerophylla. We name and describe Boana platanera sp. nov. from the southern versant of the Cordillera de Mérida (08º48'26'' N, 70º30'46'' W, WGS 84; 947 m asl), Venezuela, and refer all the populations north of the Orinoco River currently identified as B. xerophylla to this species. The new species can be readily diagnosed from B. xerophylla (characters of the latter in parentheses) by a pale orange-yellow or light brown dorsal coloration (dark brown to green), palpebral membrane with dark pigments (pigments absent); pericloacal region dark brown (cream), advertisement call with shorter first note length than B. xerophylla. This study represents an empirical example regarding false negatives behind genetic thresholds for species discovery, appraising the use of integrative taxonomic approaches.

Amphibians in Zootaxa: 20 years documenting the global diversity of frogs, salamanders, and caecilians.

Zootaxa is a mega-journal that since its inception, 20 years ago, has contributed to the documentation of the planet's biodiversity. Its role concerning terrestrial vertebrates has been crucial especially for amphibians, which are the most threatened class of vertebrates. As current editors of the Amphibia section, we reviewed the state of knowledge of taxonomic publications on amphibians over the last two decades (from 2001 to 2020). Our review reveals that 2,533 frogs, 259 salamanders, and 55 caecilians have been named in these 20 years, mainly in the tropical regions of South America, Asia, and Africa. More than half (57%) of these species descriptions were published in only 10 journals. At least 827 species of the new amphibians (29% of the total) were described in Zootaxa. This mega-journal has served also as a place of publication for monographs and systematic reviews, in addition to short articles documenting the vocalizations of anurans and the morphology of embryos and larvae. Its efficient evaluation process, the freedom of manuscript length, including full-color figures, and free of cost for the authors, has made Zootaxa a favorite for amphibian researchers. In an era of accelerating rates of biodiversity loss, documenting, describing, naming, and proposing evolutionary scenarios for species is, more than ever, an urgent task.

Discovery of an additional piece of the large gymnophthalmid puzzle: a new genus and species of stream spiny lizard (Squamata: Gymnophthalmidae: Cercosaurinae) from the western Guiana Shield in Venezuela.

Gymnophthalmids are a highly diverse group of Neotropical lizards and its species richness is still in process of discovery. The incorporation of molecular evidence and a noticeable increase in taxon and geographic sampling in systematic studies has led to the description of numerous new genera and species of gymnophthalmids (particularly in Cercosaurinae) in recent years. Herein we describe a new genus and species of cercosaurine lizard with crocodile-like morphology, from the Venezuelan Guiana Shield on the basis of molecular phylogenetic and morphological evidence. Kataphraktosaurus gen. nov. can be readily distinguished from all other genera of Cercosaurinae by a unique combination of morphological characters that includes heterogeneous dorsal scalation with enlarged and strongly keeled scales forming two paravertebral rows, ventral and subcaudal scales imbricated and strongly keeled, large and symmetrical cephalic scales, absence of postmental scale, palpebral disc translucent and divided, tail slightly compressed, all digits clawed, and only six femoral pores (three at each hindlimb) inserted in a clump of small scales. This genus is described as monotypic and only contains Kataphraktosaurus ungerhamiltoni sp. nov., which is known from one specimen and diagnosed by the same set of aforementioned characters. The secretive habits of this species and the remoteness of the locality may explain its singleton situation. Following the International Union for Conservation of Nature's criteria, we categorized the new species as Data Deficient.

Redescription and phylogenetics of emAllobates/em emtrilineatus/em (Boulenger 1884 "1883") (Anura: Aromobatidae) based on topotypic specimens.

Allobates trilineatus is the second most geographically widespread species in the genus Allobates, its range extending from northern Ecuador to southern Peru along the Andean foothills of Amazonia and to the east, into Acre, Brazil. However, detailed phenotypic and genetic variation from topotypic specimens is lacking, raising doubts about the identification of specimens in the literature. To solve this problem, we collected 16 topotypic specimens-including male and female adults and juveniles-and associated data such as advertisement calls and tissue samples. Based upon this material, we redescribe the phenotypic variation within A. trilineatus and evaluate its phylogenetic position using a fragment of the mitochondrial gene 16S rDNA. Allobates trilineatus is distinguished from its congeners by its small body size (adult snout-to-vent-length = 14.6-16.6 mm), preserved males with dark gray throat, and gray chest and belly, pale dorsolateral stripe straight and conspicuous, and advertisement call formed by the emission of groups of note-pairs with dominant frequency at 5.06-5.81 kHz. Our phylogenetic results indicate that none of the specimens assigned to this species in previous phylogenetic studies cluster within the clade formed by topotypic samples, except for the sample of one tadpole. Furthermore, our comparison of published phenotypic and genetic data assigned to A. trilineatus with our new data led us to conclude that A. trilineatus as previously recognized was actually a complex of cryptic, closely related species. Although with the data at hand we cannot fully resolve the taxonomy of all sampled populations in previous studies, we provide a new definition and delimitation of A. trilineatus sensu stricto, assign other specimens to different evolutionary units corresponding to candidate species, and flag other important taxonomic issues.

Evidence of absence treated as absence of evidence: The effects of variation in the number and distribution of gaps treated as missing data on the results of standard maximum likelihood analysis.

Although numerous studies have demonstrated the theoretical and empirical importance of treating gaps as insertion/deletion (indel) events in phylogenetic analyses, the standard approach to maximum likelihood (ML) analysis employed in the vast majority of empirical studies codes gaps as nucleotides of unknown identity ("missing data"). Therefore, it is imperative to understand the empirical consequences of different numbers and distributions of gaps treated as missing data. We evaluated the effects of variation in the number and distribution of gaps (i.e., no base, coded as IUPAC "." or "-") treated as missing data (i.e., any base, coded as "?" or IUPAC "N") in standard ML analysis. We obtained alignments with variable numbers and arrangements of gaps by aligning seven diverse empirical datasets under different gap opening costs using MAFFT. We selected the optimal substitution model for each alignment using the corrected Akaike Information Criterion in jModelTest2 and searched for optimal trees using GARLI. We also employed a Monte Carlo approach to randomly replace nucleotides with gaps (treated as missing data) in an empirical dataset to understand more precisely the effects of varying their number and distribution. To compare alignments, we developed four new indices and used several existing measures to quantify the number and distribution of gaps in all alignments. Our most important finding is that ML scores correlate negatively with gap opening costs and the amount of missing data. However, this negative relationship is not due to the increase in missing data per se-which increases ML scores-but instead to the effect of gaps on nucleotide homology. These variables also cause significant but largely unpredictable effects on tree topology.

The advertisement call of Chiasmocleis (Syncope) carvalhoi (Nelson, 1975) (Anura: Microhylidae), with taxonomic remarks based on new morphological observations of specimens from Peruvian Amazonia.

We quantitatively describe for the first time the advertisement call of Chiasmocleis (Syncope) carvalhoi using recordings  of five specimens from northern Peruvian Amazonia. The advertisement call is characterized by a single, short, tonal, and high-pitched note. Call duration ranges between 0.03-0.06 s and its dominant frequency between 7.12-7.92 kHz. The lack of pulses within notes distinguishes C. (Syncope) carvalhoi from all congeneric species except C. (Syncope) antenori, C. (Syncope) parkeri and C. (Chiasmocleis) mantiqueira, which can be distinguished by the shorter duration of their notes and silent intervals (in C. (Syncope) antenori and C. (Syncope) parkeri) and by the lower dominant frequency (in C. (Syncope) parkeri and C. (Chiasmocleis) mantiqueira). In addition, based on our collected specimens, we report on and discuss about the variation of some external morphological characters and natural history of C. (Syncope) carvalhoi, including the absence of vocal slits and sacs. Our study reveals a potential association between the absence of vocal slits and sacs, and tonal calls in Chiasmocleis, as well as conflicting morphological diagnostics characters in the literature.

A ketocarotenoid-based colour polymorphism in the Sira poison frog Ranitomeya sirensis indicates novel gene interactions underlying aposematic signal variation.

The accumulation of red ketocarotenoids is an important component of coloration in many organisms, but the underlying mechanisms are poorly understood. In some organisms, ketocarotenoids are sequestered from the diet and can accumulate when enzymes responsible for carotenoid breakdown are disrupted. In other organisms, ketocarotenoids are formed endogenously from dietary precursors via oxidation reactions carried out by carotenoid ketolase enzymes. Here, we study the genetic basis of carotenoid coloration in an amphibian. We demonstrate that a red/yellow polymorphism in the dendrobatid poison frog Ranitomeya sirensis is due to the presence/absence of ketocarotenoids. Using whole-transcriptome sequencing of skins and livers, we found that a transcript encoding a cytochrome P450 enzyme (CYP3A80) is expressed 3.4-fold higher in livers of red frogs versus yellow. As CYP3A enzymes are known carotenoid ketolases in other organisms, our results point to CYP3A80 as a strong candidate for a carotenoid ketolase in amphibians. Furthermore, in red frogs, the transcript encoding the carotenoid cleavage enzyme BCO2 is expressed at a low level or as a splice variant lacking key catalytic amino acids. This suggests that BCO2 function may be disrupted in red frogs, providing a mechanism whereby the accumulation of ketocarotenoids and their dietary precursors may be enhanced.

Mechanisms for Color Convergence in a Mimetic Radiation of Poison Frogs.

In animals, bright colors often evolve to mimic other species when a resemblance is selectively favored. Understanding the proximate mechanisms underlying such color mimicry can give insights into how mimicry evolves-for example, whether color convergence evolves from a shared set of mechanisms or through the evolution of novel color production mechanisms. We studied color production mechanisms in poison frogs (Dendrobatidae), focusing on the mimicry complex of Ranitomeya imitator. Using reflectance spectrometry, skin pigment analysis, electron microscopy, and color modeling, we found that the bright colors of these frogs, both within and outside the mimicry complex, are largely structural and produced by iridophores but that color production depends crucially on interactions with pigments. Color variation and mimicry are regulated predominantly by iridophore platelet thickness and, to a lesser extent, concentration of the red pteridine pigment drosopterin. Compared with each of the four morphs of model species that it resembles, R. imitator displays greater variation in both structural and pigmentary mechanisms, which may have facilitated phenotypic divergence in this species. Analyses of nonmimetic dendrobatids in other genera demonstrate that these mechanisms are widespread within the family and that poison frogs share a complex physiological "color palette" that can produce diverse and highly reflective colors.

Vastly underestimated species richness of Amazonian salamanders (Plethodontidae: Bolitoglossa) and implications about plethodontid diversification.

We present data showing that the number of salamander species in Amazonia is vastly underestimated. We used DNA sequences of up to five genes (3 mitochondrial and 2 nuclear) of 366 specimens, 189 corresponding to 89 non-Amazonian nominal species and 177 Amazonian specimens, including types or topotypes, of eight of the nine recognized species in the region. By including representatives of all known species of Amazonian Bolitoglossa, except for one, and 73% of the currently 132 recognized species of the genus, our dataset represents the broadest sample of Bolitoglossa species, specimens, and geographic localities studied to date. We performed phylogenetic analyses using parsimony with tree-alignment and maximum likelihood (ML) with similarity alignment, with indels as binary characters. Our optimal topologies were used to delimit lineages that we assigned to nominal species and candidate new species following criteria that maximize the consilience of the current species taxonomy, monophyly, gaps in branch lengths, genetic distances, and geographic distribution. We contrasted the results of our species-delimitation protocol with those of Automated Barcode Gap Discovery (ABGD) and multi-rate Poisson Tree Processes (mPTP). Finally, we inferred the historical biogeography of South American salamanders by dating the trees and using dispersal-vicariance analysis (DIVA). Our results revealed a clade including almost all Amazonian salamanders, with a topology incompatible with just the currently recognized nine species. Following our species-delimitation criteria, we identified 44 putative species in Amazonia. Both ABGD and mPTP inferred more species than currently recognized, but their numbers (23-49) and limits vary. Our biogeographic analysis suggested a stepping-stone colonization of the Amazonian lowlands from Central America through the Chocó and the Andes, with several late dispersals from Amazonia back into the Andes. These biogeographic events are temporally concordant with an early land bridge between Central and South America (~10-15 MYA) and major landscape changes in Amazonia during the late Miocene and Pliocene, such as the drainage of the Pebas system, the establishment of the Amazon River, and the major orogeny of the northern Andes.

Neotropical frogs and mating songs: The evolution of advertisement calls in glassfrogs.

Anurans emit advertisement calls with the purpose of attracting mates and repelling conspecific competitors. The evolution of call traits is expected to be associated with the evolution of anatomical and behavioural traits due to the physics of call emission and transmission. The evolution of vocalizations might imply trade-offs with other energetically costly behaviours, such as parental care. Here, we investigated the association between body size, calling site, parental care and call properties (call duration, number of notes, peak frequency, frequency bandwidth and call structure) of the advertisement calls of glassfrogs (Centrolenidae)-a family of Neotropical, leaf-dwelling anurans-using phylogenetic comparative methods. We also explored the tempo and mode of evolution of these traits and compared them with those of three morphological traits associated with body size, locomotion and feeding. We generated and compiled acoustic data for 72 glassfrog species (46% of total species richness), including representatives of all genera. We found that almost all acoustic traits have significant, but generally modest, phylogenetic signal. Peak frequency of calls is significantly associated with body size, whereas call structure is significantly associated with calling site and paternal care. Thus, the evolution of body size, calling site and paternal care could constrain call evolution. The estimated disparity of acoustic traits was larger than that of morphological traits and the peak in disparity of acoustic traits generally occurred later in the evolution of glassfrogs, indicating a historically recent outset of the acoustic divergence in this clade.

The tadpole of Boana stellae (Anura: Hylidae).

Boana stellae (Kwet, 2008) was allocated within the B. pulchella species group and, presumably, is closely related to B. caipora (Antunes, Faivovich, and Haddad, 2008), B. curupi (Garcia, Faivovich, and Haddad, 2007), B. joaquini (Lutz, 1968), and B. semiguttata (Lutz, 1925) due to the similarity of the external morphology of adult specimens (Kwet 2008). Boana stellae is currently only known from adult specimens and embryos from the type locality in Rio Grande do Sul, southeastern Brazil, and from a single locality in Misiones, Argentina (Ferro et al. 2018). Although the embryo morphology has been described in detail (Navarro Acosta et al. 2017; Navarro Acosta Vera Candioti 2017, as Hypsiboas sp. gr. pulchellus), the tadpole of B. stellae has not been yet described.

The taxonomic status of Petropedetes newtonii (Amphibia, Anura, Petropedetidae).

The taxon Petropedetes newtonii was described in 1895 by Bocage, from Bioko Island (Equatorial Guinea). This taxon, whose holotype is lost, has been misidentified since Boulenger's revision of the genus in 1900 and its relationships with other taxa (P. vulpiae and P. johnstoni) is confusing. Currently, P. newtonii is considered a synonym of P. johnstoni. In this work, by revising morphological characters of non-webbed Petropedetes of Bioko, we demonstrate the morphological singularity of these specimens with respect to P. johnstoni and P. vulpiae and their association with the name Petropedetes newtonii. Consequently, we provide the subsequent designation of a neotype of P. newtonii and revalidate this species from its synonym with P. johnstoni.