Search

The Neotropical genus Scinax Wagler currently comprises 127 species of small treefrogs distributed from southern Mexico to Argentina and Uruguay, including some islands such as Trinidad and Tobago, and St. Lucia (Frost 2020). Two major clades are recognized within Scinax, the S. catharinae and the S. ruber clades. The former is composed of two species groups, the S. catharinae and the S. perpusillus groups (Faivovich 2002; Faivovich et al. 2005). The S. perpusillus species group currently comprises 13 species: S. alcatraz (Lutz); S. arduous Peixoto; S. atratus (Peixoto); S. belloni Faivovich, Gasparini Haddad; S. cosenzai Lacerda, Peixoto Feio; S. faivovichi Brasileiro, Oyamaguchi Haddad; S. insperatus Silva Alves-Silva; S. littoreus (Peixoto); S. melloi (Peixoto), S. peixotoi Brasileiro, Haddad, Sawaya Martins; S. perpusillus (Lutz Lutz); S. tupinamba Silva Alves-Silva; and S. v-signatus (Lutz). These species are endemic of the Brazilian Atlantic Forest and are recognized by their intimate association with bromeliads (Fig. 1), in which adults breed and lay their eggs, and tadpoles develop (Peixoto 1987, 1995; Alves-Silva Silva 2009).

The external larval morphology of three species of the microhylid genus Kaloula (K. indochinensis, K. mediolineata, and K. pulchra) inhabiting the southern and central regions of Vietnam is studied. The similarities and the distinctive features of their morphometric characters, the structure of their mouthparts and spiracle, and their coloration are analysed with consideration of the geographic variability. A description of the tadpole of K. indochinensis is provided for the first time. The interspecific comparison revealed the shape of the mouthparts and the spiracle as the most reliable diagnostic characters for the field identification of the coexistent Kaloula tadpoles. The first description of the larval chondrocranium and hyobranchial apparatus of K. indochinensis demonstrates a set of morphological traits characteristic of suspension-feeding microhylids. Some developmental parameters (egg number and size, duration of embryonic and larval development, larvae size and stage at hatching) are provided for K. indochinensis and K. pulchra.

Fauna of the Central Ryukyus includes a high percentage of endemic species, and Hallowell's tree frog Hyla hallowellii Thompson, 1912 is one of such elements, occurring in a total of eight islands in the Amami and Okinawa Island groups. Using samples representing all of these eight island populations, we studied variations in morphology, karyotype, allozyme, and mtDNA, to clarify the pattern of geographic differentiation of H. hallowellii and consider factors for its formation. We could not clearly discriminate one population from another in morphology, nor could we find any interpopulation difference in karyotype. From genetic analyses, using allozymes and cyt b , we found low overall differentiations among populations. However, the southern populations from Okinawajima and Yoronjima were genetically nearly identical with the northern Amamioshima population. From that group the geographically intermediate Tokunoshima and Kakeromajima populations showed prominent differentiations. These patterns of geographical differentiation greatly differ from those known in other amphibian species of the Central Ryukyus, and suggest that H. hallowellii has evolutionary history unique to the species.

The liver is an important metabolic organ in vertebrates. In anurans, the hepatosomatic index (HSI) reflects differences in energy storage and reproductive activities between males and females. The objective of this study was to describe the histological and histometric parameters of the livers of five species of Neotropical anurans, taking sex-related differences into account. We also tested how the relationship between quantitative histometric variables and HSI varied between males and females in different species. Five males and five females of Elachistocleis matogrosso, Leptodactylus podicipinus, Lysapsus limellum, Pseudis platensis, and Trachycephalus typhonius were captured in central Brazil during the rainy season. HSI did not vary according to sex, but it varied among species. Elachistocleis matogrosso had the highest HSI due to the large hepatocyte size. The percentage of melanomacrophage centers (MMCs) was higher in P. platensis and L. limellum. In T. thyphonius, hepatocyte area was negatively associated with HSI, while the MMC percentages were positively associated with HSI. The liver plays a key role in reproductive activities, especially for species with explosive reproduction. Additionally, histometric patterns and volumetric structural density varied between males and females due to energy utilization for reproduction. Not only are these results important for future studies on hepatic morphophysiology but they also provide tools for evolutionary and phylogenetic studies., (© 2020 American Association for Anatomy.)

The external morphology, internal oral morphology, the chondrocranium and the hyobranchial apparatus of Elachistocleis bumbameuboi, E. cf. piauiensis, E. cesarii and E. bicolor are described and compared with each other and with other species of microhylids using available descriptions from the literature. The general morphology of Elachistocleis species is conservative in many aspects. Differences between species are subtle and are found in the body shape, the edge of the snout, fin height, if the lateral line is evident, the presence of regular pustules in the buccal roof arena, the posterolateral edge of the cartilago suprarostralis, the shape of the fenestra in the occipital region, presence or absence of fenestra hypophysea, the margin of the processus antorbitalis, expansions in the ventrolateral process, the shape and inclination of the fenestra subocularis, whether the subotic process is single or slightly bifid, and the inclination of the processus anterolateralis hyalis., (© 2021 Wiley Periodicals LLC.)

During the Natufian period, more than 12,000 years ago, Eynan (Ain Mallaha) was an important human settlement in the Hula Valley, Israel. This study concentrates on the anuran and squamate assemblage from the ultimate stage of the Natufian period at the site, the Final Natufian. Over five thousand bones assigned to at least sixteen taxa were studied from a sampled segment of the excavated open-air site. Relative species abundance, spatial distribution, taphonomic observations and ecological considerations all pointed to the conclusion that the inhabitants of Eynan intensively exploited three large "colubrine" snakes species: the Large Whip Snake (Dolichophis jugularis), the Eastern Montpellier Snake (Malpolon insignitus) and an Eastern Four-lined Ratsnake (Elaphe cf. sauromates). These snakes were the most desired and were intensively gathered, while other snakes and lizards could have been opportunistically collected when encountered. We raise questions about whether the large "colubrines" exploitation should be interpreted as additional evidence of increasing diet breadth. We suggest challenging this line of reasoning and offer possible alternative motives., Competing Interests: The authors have declared that no competing interests exist.

Among anuran species with biphasic life cycle, the occurrence of intraspecific larval morphotypes has been related to variations in developmental time, diet, geographical variation, or response to predators. Here, we evaluated the external morphological variation of larvae among three populations, located more than 270 km apart, of the anuran hylid Scinax fuscovarius by linear and geometric methods, to elucidate the presence of geographically different morphs. Comparisons targeted development, growth, and external morphology. Studied populations exhibited differences in reproductive seasonality, growth rate, timing of development, shape, and size. Shape and size comparisons revealed two well-differentiated morphs, one of them shared by the two closest populations. Morphological differences evidenced a smaller and depressed form of the entire body plan in the most distant population, which showed continuous reproduction throughout the rainy season and under more unpredictable conditions. We interpret the occurrence of the two different larval morphs in S. fuscovarius as a by-product of local geographical conditions, and discuss on possible associations with biotic and abiotic factors cues., (Copyright © 2021 Elsevier GmbH. All rights reserved.)

As anuran saltatory locomotion has specific functional requirements achieved through certain intra- and inter-limb proportions, we analyzed pattern and degree of morphological integration in limbs of ten anuran species to reveal the relationship of shared developmental programs of serially homologous structures and locomotor specialization. Our main objectives were (1) to examine if morphological and functional differences in forelimb and hindlimb were associated with reduced covariation between limbs, (2) and to reveal patterns of correlation between species and the roles played by evolutionary history (phylogeny) and ecology (lifestyle and habitat use). Species with different locomotor behaviours (walking, jumping, hopping, running, climbing, swimming and burrowing) were used. Partial correlations showed that species shared similar patterns of functionally based morphological integration, with increased correlations in elements within limbs and reduced correlations between limbs. This was mainly based on strong correlations between proximal elements, humerus-radioulna and femur-tibiofibula. To test the influence of phylogenetic relationships and ecological demands we used different matrices (correlation similarity matrix, ecological similarity matrix, matrices of phylogenetic distance and morphological distance). The changes in correlation patterns are shown to be dissociated from phylogeny. On the other hand, they are to some extent shaped by habitat use and locomotion, as the species with similar locomotor behaviour also tend to have stronger similarity in integration patterns. The results from this study provide insight into the processes underlying the evolutionary change of anuran limbs, highlighting function as the main factor that shaped morphological integration of the examined species., (Copyright © 2020 Elsevier GmbH. All rights reserved.)

The origin of morphological diversity is a critical question in evolutionary biology. Interactions between the environment and developmental processes have determining roles in morphological diversity, creating patterns through space and over time. Also, the shape of organisms tends to vary with increasing size as a result of those developmental processes, known as allometry. Several studies have demonstrated that the body sizes of anurans are associated with hydric conditions in their environments and that localities with high water stress tend to select for larger individuals. However, how environmental conditions alter those patterns of covariance between size and shape is still elusive. We used 3D geometric morphometric analyses, associated with phylogenetic comparative methods, to determine if the morphological variations and allometric patterns found in Arboranae (Anura) is linked to water conservation mechanisms. We found effects of the hydric stress on the shape of Arboranae species, favouring globular shapes. Also, the allometric patterns varied in intensity according to the water stress gradient, being particularly relevant for smaller frogs, and more intense in environments with higher water deficits. Our study provides empirical evidence that more spherical body shapes, especially among smaller species, reflect an important adaptation of anurans to water conservation in water-constrained environments.

The genus Edalorhina consists of two species of small forest-floor frogs inhabiting the Amazon basin. The tadpole of Edalorhina perezi, the most widely distributed species, was previously described based on a single and early stage (Gosner 25) individual. Herein, we provide a description of the tadpole in Gosner stages 35-36 including internal morphology data (i.e., buccopharyngeal cavity and larval skeleton) based on samples from two populations from Ecuador. Edalorhina shares a generalized morphology with most members of its closely related taxa; however, it is distinguished from the other species by having an almost terminal oral disc. The presence of a dextral vent tube is considered a synapomorphy for the clade consisting of Edalorhina, Engystomops, and Physalaemus. Within this clade, the combination of two lingual papillae, a filiform median ridge, and the lack of buccal roof papillae are diagnostic of E. perezi and putative autapomorphies of Edalorhina. Chondrocranial anatomy provides characteristics, that is, presence of and uniquely shaped processus pseudopterygoideus and cartilago suprarostralis with corpora and alae joined by dorsal and ventral connections that readily differentiates the genus from other Leiuperinae., (© 2020 Wiley Periodicals LLC.)

The axial skeleton of the anurans has undergone an evolutionary reduction of its bone elements. This structural plan is strongly preserved throughout the order and would have emerged as a highly specialized anatomical adaptation to its locomotor jumping pattern. The development programs that direct the vertebral morphogenesis of the anurans are poorly described and the molecular bases that have caused their pattern to differ from other tetrapods are completely unknown. In this work, we review the ontogeny of the spinal column of the anurans and explore the genetic mechanisms that could explain the morphological difference and the maintenance of the body plan during evolution. Here, we propose that the absence of caudal osseous elements, as a consequence of the inability of sclerotomes to form cartilaginous condensations in frogs, could be due to changes in both pattern and expression levels of Hox , Pax1 , Pax9 and Uncx4.1 genes along the anteroposterior axis. The anteriorised expression of the Hox genes together with the reduction in the expression levels of Pax1 , Pax9 and Uncx4 in the posterior somites could explain, at least partly, the loss of caudal vertebrae in the anurans during evolution.

There are remarkable interspecific differences in the sizes of the larvae of Andean frogs of the genus Telmatobius. This size variation seems to be associated with the duration of the larval stage and may affect the hind-limb morphology in Telmatobius. Larval, juvenile, and adult Telmatobius rubigo and T. oxycephalus were examined to determine the variation in relative sizes of hind-limb elements, their growth patterns during postmetamorphic life, and skeletal ontogeny. The results showed that the proportionately shorter hind limbs of T. rubigo relative to those of T. oxycephalus are associated with the protracted development and ossification of hind limbs during the prolonged larval life of T. rubigo. Postmetamorphically, the hind limbs grew faster than the body in juveniles of both species in contrast to the relative growth rates of the hind limbs and bodies of the adults. The growth phase of juvenile T. rubigo seems shorter than that of juvenile T. oxycephalus; possibly, this heightens the difference in the relative lengths of hind limbs after metamorphosis. Temperature affects the effects of thyroid hormone on growth and development, and T. rubigo lives at much higher, colder elevations than does T. oxycephalus. It is not clear whether the developmental differences described here are plastic (i.e., environmentally induced) or genetically fixed in each species., (© 2020 Wiley Periodicals LLC.)

Anurans (frogs and toads) have a unique pelvic and hind limb skeleton among tetrapods. Although their distinct body plan is primarily associated with saltation, anuran species vary in their primary locomotor mode (e.g., walkers, hoppers, jumpers, and swimmers) and are found in a wide array of microhabitats (e.g., burrowing, terrestrial, arboreal, and aquatic) with varying functional demands. Given their largely conserved body plan, morphological adaptation to these diverse niches likely results from more fine-scale morphological change. Our study determines how shape differences in Anura's unique pelvic and hind limb skeletal structures vary with microhabitat, locomotor mode, and jumping ability. Using microCT scans of preserved specimens from museum collections, we added 3D landmarks to the pelvic and hind limb skeleton of 230 anuran species. In addition, we compiled microhabitat and locomotor data from the literature for these species that span 52 of the 55 families of frogs and ∼210 million years of anuran evolution. Using this robust dataset, we examine the relationship between pelvic and hind limb morphology and phylogenetic history, allometry, microhabitat, and locomotor mode. We find pelvic and hind limb changes associated with shifts in microhabitat ("ecomorphs") and locomotor mode ("locomorphs") and directly relate those morphological changes to the jumping ability of individual species. We also reveal how individual bones vary in evolutionary rate and their association with phylogeny, body size, microhabitat, and locomotor mode. Our findings uncover previously undocumented morphological variation related to anuran ecological and locomotor diversification and link that variation to differences in jumping ability among species., (© The Author(s) 2020. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved. For permissions please email: journals.permissions@oup.com.)

Surprisingly, little is known about body-size evolution within the most diverse amphibian order, anurans (frogs and toads), despite known effects of body size on the physiological, ecological and life-history traits of animals more generally. Here, we examined anuran body-size evolution among 2,434 species with over 200 million years of shared evolutionary history. We found clade-specific evolutionary shifts to new body-size optima along with numerous independent transitions to gigantic and miniature body sizes, despite the upper limits of anuran body size remaining quite consistent throughout the fossil record. We found a weak, positive correlation between a species' body size and maximum latitude and elevation, including a dearth of small species at higher elevations and broader latitudinal and elevational ranges in larger anurans. Although we found modest differences in mean anuran body size among microhabitats, there was extensive overlap in the range of body sizes across microhabitats. Finally, we found that larger anurans are more likely to consume vertebrate prey than smaller anurans are and that species with a free-swimming larval phase during development are larger on average than those in which development into a froglet occurs within the egg. Overall, anuran body size does not conform to geographic and ecological patterns observed in other tetrapods but is perhaps more notable for variation in body size within geographic regions, ecologies and life histories. Here, we document this variation and propose target clades for detailed studies aimed at disentangling how and why variation in body size was generated and is maintained in anurans., (© 2020 European Society For Evolutionary Biology. Journal of Evolutionary Biology © 2020 European Society For Evolutionary Biology.)

In animal communication, receivers benefit from signals providing reliable information on signalers' traits of interest. Individuals involved in conflicts, such as competition between rivals, should pay particular attention to cues that are "unfakeable" by the senders due to the intrinsic properties of the production process. In bioacoustics, the best-known example of such "index signals" is the relationship between a sender's body size and the dominant frequency of their vocalizations. Dominant frequency may, however, not only depend on an animal's morphology but also on the interaction between the sound production system and its immediate environment. Here, we experimentally altered the environment surrounding calling frogs and assessed its impact on the signal produced. Our results show that frogs that are floating are able to inflate their vocal sacs fully and that this change in inflation level is correlated with a decrease of call dominant frequency.

David and Karin Pfennig introduce character displacement, the divergent evolution of traits in overlapping species., (Copyright © 2020 Elsevier Inc. All rights reserved.)

Abra de Zamora is an important biodiversity hotspot in southern Ecuador. Between 1938 and 2010, eleven species of frogs were described from here: Lynchius flavomaculatus, Gastrotheca psychrophila, Pristimantis balionotus, P. colodactylus, P. cryptomelas, P. percultus, P. versicolor, P. vidua, Telmatobius cirrhacelis, P. andinognomus, and Atelopus podocarpus. Unfortunately, many of these species were not re-encountered after their original description, and for the majority DNA samples were not available, making their phylogenetic position unknown. In this study, we assess the current state of the amphibians from Abra de Zamora, by: i. redescribing the species which were first reported from the area, by contributing genetic delimitation (for L. flavomaculatus, P. balionotus, P. colodactylus, P. percultus, and P. vidua), release call (L. flavomaculatus) and advertisement call descriptions (for P. balionotus, P. vidua and P. versicolor); ii. presenting an updated amphibian species list of Abra de Zamora, with the description of two additional Pristimantis species; iii. updating the distribution of these species, including data collected in similar montane habitats from surrounding areas; and iv. amending recommendations regarding their conservation status., Competing Interests: The authors have declared that no competing interests exist.

Tetrapod limbs have been used as a model system to investigate how selective pressures and constraints shape morphological evolution. Anurans have had many independent transitions to various microhabitats, allowing us to dissect how these factors influence limb morphology. Furthermore, anurans provide a unique system to test the generality of developmental constraints proposed in mammals, namely that later-developing limb bones are under less constraint and show more variation. We used microcomputed tomography scans of 236 species from 52 of 55 families, geometric morphometrics, and modern phylogenetic comparative methods to examine how limb bones are related to microhabitat, phylogeny, allometry, and developmental timing. Although there was significant phylogenetic signal, anuran limb shape showed a relationship with microhabitat and to a lesser extent, body size. We found that distal bones had higher evolutionary rates than proximal bones, providing evidence that developmental constraints are reduced in later-developing bones. Distal bones also showed increased selection related to allometry and microhabitat, providing an additional explanation for higher evolutionary rates. By looking at the evolution of limb shape across a diverse clade, we demonstrated that multiple factors have shaped anuran limbs and that greater evolutionary lability in later-developing limb bones is likely a general trend among tetrapods., (© 2020 The Authors. Evolution © 2020 The Society for the Study of Evolution.)

The free living larvae of anurans (i.e., tadpoles) are a key element in the evolution and diversification of this group, and as such, their morphology is an important element to understand the phylogenetic relationships of frogs. However, the lack of data on larval morphology prevents us from fully understanding larval evolution in several lineages. The Neotropical genus Proceratophrys currently comprises 39 species, but descriptions of the internal morphology of larvae in this group are rare and restricted to few aspects of their buccopharyngeal cavity, chondrocranium, and muscles. In the present study, I describe the internal anatomy of the tadpole of P. minuta and report a new remarkable myological character state for the species. Given the rarity of this material, the description of this species' buccopharyngeal and musculo-skeletal elements is based on two tadpoles in developmental stages 30 and 31. Several new apomorphic character states are described: (a) the presence of a conical papilla in the interior of the nostril; (b) a row of five short, conical papilla preceding the tall, postnarial papilla; and (c) the m. mandibulolabialis inserting in the gular skin. This latter feature is a remarkable, newly discovered character state that had never been reported in the literature before and is probably related to a particular feeding habit of the tadpoles of this species. The function of the m. mandibulolabialis in P. minuta is unknown., (© 2020 Wiley Periodicals LLC.)

Species loss by habitat replacement operating as an ecological filter is a well-known consequence of modern human activities. In contrast, the ecological and evolutionary response of species overcoming those filters in converted habitats has not been thoroughly explored. Species that persist are subject to novel and potentially stressful conditions that may induce certain morphological changes. We evaluated changes in the functional morphology of three anuran species persisting after the conversion of areas of the Atlantic Forest into pine plantations. We specifically evaluated differences in body size and body condition indices, head width, and hind legs' length between adult individuals from both habitats and sexes. Habitat conversion and sexual dimorphism affected the morphology of the three anurans, with varying effects upon species and traits. Regarding the effect of habitat conversion, Elachistocleis bicolor increased body condition in plantations with no changes in the other traits, Physalaemus cuvieri showed only a marginal increment in residual body mass in plantations, and Odontophrynus americanus exhibited a substantial increment in body size while maintaining its body condition in plantations. Remarkably, none of the results suggested these persistent anurans were stressed by forest conversion. This study shows that habitat conversion may induce intraspecific morphological changes in ecologically relevant traits of persistent species, and that disturbed areas do not necessarily imply stressful, low quality habitats affecting all persistent native species negatively., (© 2020 International Society of Zoological Sciences, Institute of Zoology/Chinese Academy of Sciences and John Wiley & Sons Australia, Ltd.)

The majority of animal species have complex life cycles, in which larval stages may have very different morphologies and ecologies relative to adults. Anurans (frogs) provide a particularly striking example. However, the extent to which larval and adult morphologies (e.g. body size) are correlated among species has not been broadly tested in any major group. Recent studies have suggested that larval and adult morphology are evolutionarily decoupled in frogs, but focused within families and did not compare the evolution of body sizes. Here, we test for correlated evolution of adult and larval body size across 542 species from 42 families, including most families with a tadpole stage. We find strong phylogenetic signal in larval and adult body sizes, and find that both traits are significantly and positively related across frogs. However, this relationship varies dramatically among clades, from strongly positive to weakly negative. Furthermore, rates of evolution for both variables are largely decoupled among clades. Thus, some clades have high rates of adult body-size evolution but low rates in tadpole body size (and vice versa). Overall, we show for the first time that body sizes are generally related between adult and larval stages across a major group, even as evolutionary rates of larval and adult size are largely decoupled among species and clades.

Competition over mates is a powerful force shaping trait evolution. For instance, better cognitive abilities may be beneficial in male-male competition and thus be selected for by intrasexual selection. Alternatively, investment in physical attributes favoring male performance in competition for mates may lower the resources available for brain development, and more intense male mate competition would coincide with smaller brains. To date, only indirect evidence for such relationships exists, and most studies are heavily biased toward primates and other homoeothermic vertebrates. We tested the association between male brain size (relative to body size) and male-male competition across N = 30 species of Chinese anurans. Three indicators of the intensity of male mate competition-operational sex ratio (OSR), spawning-site density, and male forelimb muscle mass-were positively associated with relative brain size, whereas the absolute spawning group size was not. The relationship with the OSR and male forelimb muscle mass was stronger for the male than for the female brains. Taken together, our findings suggest that the increased cognitive abilities of larger brains are beneficial in male-male competition. This study adds taxonomic breadth to the mounting evidence for a prominent role of sexual selection in vertebrate brain evolution.

The current functional model of the anuran larynx includes four pairs of laryngeal muscles. Their contractions do not account, however, for the behavioral control of call complexity observed in male túngara frogs (Engystomops pustulosus), which optionally add a secondary note with distinct harmonic structure to their advertisement call. Examination of the túngara frog's laryngeal morphology through dissection and resin histology has revealed that the m. dilatator laryngis is divided into two separate bundles (superficial and deep). The superficial bundle closely matches the typical description of the m. dilatator laryngis and is well positioned to open the glottis. The deep bundle is exclusively innervated by the short laryngeal nerve and has an attachment to the fibrous mass, an internal laryngeal structure necessary for complex call production. This attachment indicates a separate role for the deep bundle in controlling the complexity of the call. Based on physical separation, exclusive attachments, distinct fiber orientation, exclusive innervation, and potential action, we recognize the deep bundle of the m. dilatator laryngis as a separate muscle. We also revalidate the name m. arylabialis which had been previously used to describe it. The split of the m. dilatator laryngis into two muscles results in a laryngeal innervation pattern that closely matches that of mammals. This study identified a novel laryngeal muscle in túngara frogs, a potential mechanism for the control of call complexity, and revealed new evidence of homologies between the laryngeal structures of amphibians and mammals. Anat Rec, 2019. © 2019 American Association for Anatomy Anat Rec, 303:1966-1976, 2020. © 2019 American Association for Anatomy., (© 2019 American Association for Anatomy.)

Species can evolve diverse strategies to survive periods of uncertainty. Animals may either invest in energy storage, allowing them to decrease foraging costs, such as locomotion or risk of predation, or they may invest in better cognitive abilities helping them to flexibly adapt their behavior to meet novel challenges. Here, we test this idea of a fat-brain trade-off in 38 species of Chinese anurans by relating the coefficient of variation of body mass (CV bodymass ; as an indicator of how much animals invest into storage over the season) to brain anatomical features. After correcting for shared ancestry and body mass, we found a negative relationship between relative brain size and CV bodymass . This indicates that anurans seem to trade-off physiological and cognitive buffering during energy shortages. As similar patterns have been reported in arboreal mammals and primates our findings suggest that the fat-brain trade-off, where animals either invest into physiological or cognitive strategies to survive harsh conditions, may be a general pattern across vertebrates., (© 2020 The Authors. Evolution © 2020 The Society for the Study of Evolution.)

Glyphosate-based herbicide (GBH) using is increasing on a global scale. Few studies have investigated the sub-lethal effects of GBH in endemic amphibian species. The present work tested the GBH Roundup Active® on the tadpoles of Dendropsophus molitor. The exposure was in a range of plausible environmental concentrations (0-0.75 μg a.e./L) during a month. D. molitor is an endemic tropical frog of South America. The exposure from 325 μg a.e./L caused histological alterations in the liver. The high-resolution optical microscopy (HROM) detected sinusoidal dilatation and cytoplasmic vacuolization. The transmission electron microscopy (TEM) showed disorganization of the endoplasmic reticulum. Since the liver is essential for detoxification, these results suggest choric effects. Exposure to another GBH has caused histological alterations in liver tadpoles liver in a previous study, but, this study tested another endemic South-American frog for only 96h. The present work applied HROM to observe lipid alterations since it does not use organic solvents; and TEM for the ultrastructural observation of hepatocytes. Environmental risk of GBH can improve by including sub-lethal effects in endemic species., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

While comparatively few amphibian species have been described on the North East Asian mainland in the last decades, several species have been the subject of taxonomical debates in relation to the Yellow sea. Here, we sampled Dryophytes sp. treefrogs from the Republic of Korea, the Democratic People's Republic of Korea and the People's Republic of China to clarify the status of this clade around the Yellow sea and determine the impact of sea level change on treefrogs' phylogenetic relationships. Based on genetics, call properties, adult morphology, tadpole morphology and niche modelling, we determined the segregated status species of D. suweonensis and D. immaculatus. We then proceeded to describe a new treefrog species, D. flaviventris sp. nov., from the central lowlands of the Republic of Korea. The new species is geographically segregated from D. suweonensis by the Chilgap mountain range and known to occur only in the area of Buyeo, Nonsan and Iksan in the Republic of Korea. While the Yellow sea is the principal element to the current isolation of the three clades, the paleorivers of the Yellow sea basin are likely to have been the major factor for the divergences within this clade. We recommend conducting rapid conservation assessments as these species are present on very narrow and declining ranges., Competing Interests: The authors have declared that no competing interests exist.

Evolutionary integration (covariation) of traits has long fascinated biologists because of its potential to elucidate factors that have shaped morphological evolution. Studies of tetrapod crania have identified patterns of evolutionary integration that reflect functional or developmental interactions among traits, but no studies to date have sampled widely across the species-rich lissamphibian order Anura (frogs). Frogs exhibit a vast range of cranial morphologies, life history strategies, and ecologies. Here, using high-density morphometrics we capture cranial morphology for 172 anuran species, sampling every extant family. We quantify the pattern of evolutionary modularity in the frog skull and compare patterns in taxa with different life history modes. Evolutionary changes across the anuran cranium are highly modular, with a well-integrated "suspensorium" involved in feeding. This pattern is strikingly similar to that identified for caecilian and salamander crania, suggesting replication of patterns of evolutionary integration across Lissamphibia. Surprisingly, possession of a feeding larval stage has no notable influence on cranial integration across frogs. However, late-ossifying bones exhibit higher integration than early-ossifying bones. Finally, anuran cranial modules show diverse morphological disparities, supporting the hypothesis that modular variation allows mosaic evolution of the cranium, but we find no consistent relationship between degree of within-module integration and disparity., (© 2020 The Authors. Evolution published by Wiley Periodicals LLC on behalf of The Society for the Study of Evolution.)

The origin and evolution of the vertebrate skull have been topics of intense study for more than two centuries. Whereas early theories of skull origin, such as the influential vertebral theory, have been largely refuted with respect to the anterior (pre-otic) region of the skull, the posterior (post-otic) region is known to be derived from the anteriormost paraxial segments, i.e. the somites. Here we review the morphology and development of the occiput in both living and extinct tetrapods, taking into account revised knowledge of skull development by augmenting historical accounts with recent data. When occipital composition is evaluated relative to its position along the neural axis, and specifically to the hypoglossal nerve complex, much of the apparent interspecific variation in the location of the skull-neck boundary stabilizes in a phylogenetically informative way. Based on this criterion, three distinct conditions are identified in (i) frogs, (ii) salamanders and caecilians, and (iii) amniotes. The position of the posteriormost occipital segment relative to the hypoglossal nerve is key to understanding the evolution of the posterior limit of the skull. By using cranial foramina as osteological proxies of the hypoglossal nerve, a survey of fossil taxa reveals the amniote condition to be present at the base of Tetrapoda. This result challenges traditional theories of cranial evolution, which posit translocation of the occiput to a more posterior location in amniotes relative to lissamphibians (frogs, salamanders, caecilians), and instead supports the largely overlooked hypothesis that the reduced occiput in lissamphibians is secondarily derived. Recent advances in our understanding of the genetic basis of axial patterning and its regulation in amniotes support the hypothesis that the lissamphibian occipital form may have arisen as the product of a homeotic shift in segment fate from an amniote-like condition., (© 2020 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society.)

The vertebrate head as a major novelty is directly linked to the evolutionary success of the vertebrates. Sequential information on the embryonic pattern of cartilaginous head development are scarce, but important for the understanding of its evolution. In this study, we use the oriental fire bellied toad, Bombina orientalis, a basal anuran to investigate the sequence and timing of larval cartilaginous development of the head skeleton from the appearance of mesenchymal Anlagen in post-neurulation stages until the premetamorphic larvae. We use different methodological approaches like classic histology, clearing and staining, and antibody staining to examine the larval skeletal morphology. Our results show that in contrast to other vertebrates, the ceratohyals are the first centers of chondrification. They are followed by the palatoquadrate and the basihyal. The latter later fuses to the ceratohyal and the branchial basket. Anterior elements like Meckel's cartilage and the rostralia are delayed in development and alter the ancestral anterior posterior pattern observed in other vertebrates. The ceratobranchials I-IV, components of the branchial basket, follow this strict anterior-posterior pattern of chondrification as reported in other amphibians. Chondrification of different skeletal elements follows a distinct pattern and the larval skeleton is nearly fully developed at Gosner Stage 28. We provide baseline data on the pattern and timing of early cartilage development in a basal anuran species, which may serve as guidance for further experimental studies in this species as well as an important basis for the understanding of the evolutionary changes in head development among amphibians and vertebrates., (© 2020 The Authors. Journal of Morphology published by Wiley Periodicals, Inc.)

Micryletta inornata is a complex species that is widely distributed from Sumatra to mainland Asia, including the Thai-Malay Peninsula and Indochina. Recently, this species was confirmed to be endemic to regions near the type locality in Sumatra, and the populations from other regions were suggested to be different species. We examined phenotypic and genotypic characters of the Sumatran populations and found an unnamed lineage in addition to the true M. inornata . The newly found lineage can be distinguished from M. inornata and other congeners by both molecular and morphological traits and has been named Micryletta sumatrana sp. nov. The new species is characterized by having a small body size, golden brown dorsum with scattered dark spots, dark brown ventrum with diffuse cream mottling, dark brown lateral head with cream spots on lips and the tympanum region extending to the axilla, and tibiotarsal articulation reaching to the front of the eye. We discuss the taxonomic status of so-called M. inornata occurring outside of its type locality, especially of M. inornata lineata .

Life history theory predicts that animals often produce fewer offspring of larger size and indicate a stronger trade-off between the number and size of offspring to cope with increasing environmental stress. In order to evaluate this prediction, we tested the life history characteristics of Bufo minshanicus at eight different altitudes on the eastern Tibetan Plateau, China. Our results revealed a positive correlation between female SVL and clutch size or egg size, revealing that larger females produce more and larger eggs. However, high-altitude toads seem to favor more offspring and smaller egg sizes when removing the effect of female SVL, which is counter to theoretical predictions. In addition, there was an overall significantly negative relationship between egg size and clutch size, indicative of a trade-off between egg size and fecundity. Therefore, we suggest that higher fecundity, rather than larger egg size, is a more effective reproductive strategy for this species of anuran living at high-altitude environments.

Frogs (Anura) are one of the most diverse vertebrate orders, comprising more than 7,000 species with a worldwide distribution and extensive ecological diversity. In contrast to other tetrapods, frogs have a highly derived body plan and simplified skull. In many lineages of anurans, increased mineralization has led to hyperossified skulls, but the function of this trait and its relationship with other aspects of head morphology are largely unexplored. Using three-dimensional morphological data from 158 species representing all frog families, we assessed wide-scale patterns of shape variation across all major lineages, reconstructed the evolutionary history of cranial hyperossification across the anuran phylogeny, and tested for relationships between ecology, skull shape, and hyperossification. Although many frogs share a conserved skull shape, several extreme forms have repeatedly evolved that commonly are associated with hyperossification, which has evolved independently more than 25 times. Variation in cranial shape is not explained by phylogenetic relatedness but is correlated with shifts in body size and ecology. The species with highly divergent, hyperossified skulls often have a specialized diet or a unique predator defense mechanism. Thus, the evolution of hyperossification has repeatedly facilitated the expansion of the head into multiple new shapes and functions., Competing Interests: The authors declare no competing interest., (Copyright © 2020 the Author(s). Published by PNAS.)

Based on anatomical and functional data, the habenula-a phylogenetically old brain structure present in all vertebrates-takes part in the integration of limbic, sensory, and basal ganglia information to guide effective response strategies appropriate to environmental conditions. In the present study, we investigated the connections of the habenular nuclei of the oriental fire-bellied toad, Bombina orientalis, and compared them with published data from lampreys, chondrichthyes, teleosts, reptiles, birds, and mammals. During phylogenetic development, the primordial habenula circuitry underwent various evolutionary adaptations and in the tetrapod line, the circuit complexity increased. The habenula circuitry of anuran amphibians, decedents of the first land-living tetrapods, seem to exhibit a mix of ancient as well as modern features. The anuran medial and lateral habenula homologs receive differential input from the septum, nucleus of the diagonal band of Broca, preoptic area, hypothalamus, rostral pallium, nucleus accumbens, ventral pallidum, and bed nucleus of the stria terminalis. Additional input arises from a border region in the ventral prethalamus, here discussed as a putative homolog of the entopeduncular nucleus of rodents. The habenular subnuclei also differentially innervate the interpeduncular nucleus, raphe nuclei, substantia nigra pars compacta and ventral tegmental area homologs, superficial mamillary area, laterodorsal tegmental nucleus, locus coeruleus, inferior and superior colliculus homologs, hypothalamus, preoptic area, septum, nucleus of the diagonal band of Broca, and main olfactory bulb. It seems likely that the main connectivity between the habenula and the basal ganglia, limbic, and sensory systems was already present in the common tetrapod ancestor., (© 2019 Wiley Periodicals, Inc.)

Whereas our understanding of the dopaminergic system in mammals allows for a distinction between ventral tegmental area (VTA) and substantia nigra pars compacta (SNc), no clear evidence for separate structures in anamniotes has been presented to date. To broaden the insight into the organization and regulation of neuromodulatory systems in anuran amphibians, tracing and immunohistochemical investigations were performed in the Oriental fire-bellied toad, Bombina orientalis. Topographically organized catecholaminergic "nigrostriatal," "mesolimbic," "mesocortical," and spinal cord projections arising from the posterior tubercle and mesencephalic tegmentum were identified. We compared these results with published data from lampreys, chondrichthyes, teleosts, amphibians, reptiles, birds, and mammals. Based on the pattern of organization, as well as the differential innervation by the habenular nuclei, domains gradually comparable to the mammalian paranigral VTA, ventral tier of the SNc, interfascicular nucleus of the VTA, and supramamillary/retromamillary area were identified. Additionally, we could demonstrate topographic separate populations of habenula neurons projecting via a direct excitatory or indirect GABAergic pathway onto the catecholaminergic VTA/SNc homologs and serotonergic raphe nuclei. The indirect GABAergic habenula pathway derives from neurons in the superficial mamillary area, which in terms of its connectivity and chemoarchitecture resembles the mammalian rostromedial tegmental nucleus. These results demonstrate a much more elaborate interconnection principle of the anuran dopaminergic system than previously assumed. Based on the data presented it seems that most features of the dopaminergic system of amniotes had already evolved when the amphibian line of evolution diverged from that leading up to mammals, reptiles, and birds., (© 2019 Wiley Periodicals, Inc.)

The genus-level recognition of monophyletic short-legged toads ( Brachytarsophrys ) has been recently implicated in the taxonomic debate of Megophrys sensu lato . In the present study, Brachytarsophrys is reasonably regarded as a distinct genus based on significant morphological differentiations and recent molecular analyses. Furthermore, a comprehensive review of this genus is performed, with two species groups proposed based on morphological differences and phylogenetic relationships. Particularly, Brachytarsophrys platyparietus is removed as a synonym of Brachytarsophrys carinense and considered a valid species due to significant genetic divergence and distinct morphological differences. In addition, a new species, Brachytarsophrys orientalis sp . nov ., is described based on a series of specimens collected from southeastern China. This work takes the member species of the genus Brachytarsophrys to seven, suggesting that the diversity of Brachytarsophrys is underestimated. In addition, the genus levels of other monophyletic groups within the subfamily Megophryinae are discussed.

The Pristimantis genus (Anura: Craugastoridae) is the most diverse among all vertebrates with 531 described species. The highest diversity occurs in Ecuador (215 species), followed by Colombia (202), Peru (139), Venezuela (60), Brazil (30), Bolivia (17), Guyana (6) Suriname and French Guiana (5). The genus is divided into 11 species groups. Of these, the P. conspicillatus group (containing 34 species), distributed in extreme southeastern Costa Rica, Isla Taboga (Panama), northern South America (from Colombia to eastern Guyana), south Bolivia, and is the best represented in Brazil (16 species). The main characteristics of this group are the tympanic membrane and tympanic annulus distinct (except in P. johannesdei); dorsum smooth or shagreen; dorsal lateral fold present or absent; usually smooth belly, but may be weakly granular in some species; toe V slightly larger than the toe III. Most of the taxonomic inconsistencies in species of Pristimantis could be due to its much conserved morphology and the lack of comprehensive taxonomic evaluations. Thus, an ongoing challenge for taxonomists dealing with the Pristimantis genus is the ubiquitous abundance of cryptic species. In this context, accurate species delimitation should integrate evidences of morphological, molecular, bioacoustics and ecological data, among others. Based on an integrative taxonomy perspective, we utilize morphological, molecular (mtDNA) and bioacoustic evidence to describe four new species of the Pristimantis conspicillatus group from the eastern Amazon basin. Pristimantis giorgii sp. nov. is known from the Xingu/Tocantins interfluve and can be distinguished from the other Pristimantis species of the region by presenting discoidal fold, dorsolateral fold absent, vocalization composed of three to four notes and genetic distance of 7.7% (16S) and 14.8% (COI) from P. latro, the sister and sympatric species with respect P. giorgii sp. nov.. Pristimantis pictus sp. nov. is known to the northern Mato Grosso state, Brazil, and can be distinguished from the other species of Pristimantis by presenting the posterior surface of the thigh with light yellow patches on a brown background, also extending to the inguinal region, vocalization consisting of four to five notes and a genetic distance of 11.6% (16S) and 19.7% (COI) from P. pluvian sp. nov., which occurs in sympatry. Pristimantis pluvian sp. nov. is known to the northern Mato Grosso state, Brazil, and may be distinguished from the other Pristimantis species by having a posterior surface of the thigh reddish and vocalization composed of two notes. Pristimantis moa sp. nov. is known to the northern Tocantins state and southwestern Maranhão state. This species can be distinguished from the other Pristimantis species by possessing slightly perceptible canthal stripe, external thigh surface with dark yellow spots on brown background, vocalization consisting of three to five notes and genetic distance of 2.3-11.7 (16S) and 10.5-23.1 (COI) for the new Pristimantis species of this study., Competing Interests: The authors have declared that no competing interests exist.

Isolated amphibian bones from the early Pliocene of Kanapoi (West Turkana, Kenya) help to improve the scarce fossil record of the late Neogene and Quaternary amphibians from East Africa. All currently available 579 bones are referable exclusively to the Anura (frogs and toads). More than half of the remains (366) are identified as Hemisus cf. Hemisus marmoratus, an extant species that still inhabits Kenya, but apparently not the northwest of the country and the Turkana area in particular. The rest of the remains are identified simply as Anura indet. because of poor preservation or non congruence with the relatively few African extant taxa whose osteology is known in detail. The Hemisus material represents the first fossil record for Hemisotidae, an endemic African family of peculiar, head-first burrowing frogs, whose sister taxon relationships indicate a divergence from brevicipitids in the Late Cretaceous or early Paleocene. The ecological requirements of extant H. marmoratus suggest that the Kanapoi area surrounding the fluvial and deltaic settings, from where the fossil remains of vertebrates were buried, was likely a grassland or relatively dry, open low tree-shrub savanna., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Taxonomists always have had intense discussions about how species should be delimited and recently many studies have used integrative approaches by combining molecular, morphological, and bioacoustic data. Although these studies are paramount for understanding species diversity, few of them actually formalize species delimitations to the final step of nomenclatural acts. Historically, the Neotropical frog genus Adenomera has been considered as a difficult taxonomic group because it comprises many morphologically similar species exhibiting high levels of intraspecific polymorphism. A recent work using molecular data shed light on the phylogenetic relationships within the genus and identified several lineages that may correspond to undescribed species but did not delimit species boundaries. In the Atlantic Forest, a clade formed by A. marmorata and two putative species (Adenomera sp. J and Adenomera sp. K) were identified. In this paper, we combine morphological, acoustic, and molecular data in order to evaluate species limits within this Atlantic Forest Adenomera clade. We provide a redescription of A. marmorata and restrict its type locality to the Tijuca Massif, in the Municipality of Rio de Janeiro, Brazil. Our results do not support A. marmorata and the two candidate species as diagnosable distinct species. Therefore A. marmorata corresponds to a species with pronounced morphological and acoustic variation in the genus and a complex phylogeographic structure., Competing Interests: The authors have declared that no competing interests exist.

Developmental novelties often underlie the evolutionary origins of key metazoan features. The anuran urostyle, which evolved nearly 200 MYA, is one such structure. It forms as the tail regresses during metamorphosis, when locomotion changes from an axial-driven mode in larvae to a limb-driven one in adult frogs. The urostyle comprises of a coccyx and a hypochord. The coccyx forms by fusion of caudal vertebrae and has evolved repeatedly across vertebrates. However, the contribution of an ossifying hypochord to the coccyx in anurans is unique among vertebrates and remains a developmental enigma. Here, we focus on the developmental changes that lead to the anuran urostyle, with an emphasis on understanding the ossifying hypochord. We find that the coccyx and hypochord have two different developmental histories: First, the development of the coccyx initiates before metamorphic climax whereas the ossifying hypochord undergoes rapid ossification and hypertrophy; second, thyroid hormone directly affects hypochord formation and appears to have a secondary effect on the coccygeal portion of the urostyle. The embryonic hypochord is known to play a significant role in the positioning of the dorsal aorta (DA), but the reason for hypochordal ossification remains obscure. Our results suggest that the ossifying hypochord plays a role in remodeling the DA in the newly forming adult body by partially occluding the DA in the tail. We propose that the ossifying hypochord-induced loss of the tail during metamorphosis has enabled the evolution of the unique anuran bauplan ., Competing Interests: The authors declare no competing interest., (Copyright © 2020 the Author(s). Published by PNAS.)

Horned Frogs of the family Ceratophryidae are conspicuous anurans represented by three endemic South American genera. Most ceratophryids inhabit semiarid environments, but three species of Ceratophrys occupy tropical or temperate humid areas. Several morphological and behavioral characters of larvae and adults are conserved across the family. Based on examination of specimens and accounts in the literature, the embryonic development of C. ornata, C. cranwelli, and the monotypic genus Chacophrys are described and compared with that of species of Lepidobatrachus. Ceratophryid embryos share a suite of morphological features and heterochronic shifts during development. Most features, such as gill structure, ciliation, early hatching, and precocious differentiation of the gut and hind limbs, are shared by all the species regardless the differences in the habitats that occupy. This is consistent with previous observations of some adult characters, and likely supports the hypothesis of an early diversification of ceratophryids in semiarid environments. Other embryonic features, such as the morphology and ontogeny of the oral disc and digestive tract, are correlated with larval feeding habits and vary within the family. The evolutionary and ecological significance of some conserved characters (e.g., gastrulation pattern, Type-A adhesive glands) and other taxon-specific features (e.g., nasal appendix) remain to be explored in the group., (© 2019 Wiley Periodicals, Inc.)

Several hormones regulate anuran larval development, most notably thyroid hormones (THs). In anurans, metamorphosis fails when the thyroid gland is absent or inactivated, resulting in giant tadpoles. Larval gigantism occurs naturally in neotropical frogs of the genus Pseudis as a result of a prolonged larval period. Its thyroid function is poorly investigated and the focus of this study. We describe qualitative and quantitative variations in larval development for field-captured specimens of Pseudis platensis and compare those to the development of two sympatric species, Phyllomedusa sauvagii and Pithecopus azureus, which have small tadpoles and a shorter larval period. We describe morphological changes in the thyroid glands of larval and adult specimens. In contrast to other species with similar ecological requirements, P. platensis exhibits distinct glandular activity. During premetamorphosis, there was little or no thyroid activity, a period in which the tadpole reached 70% of its maximum size. Development and degree of activity of the thyroid gland determine the duration of the early stages of the larval period. Thyroid gland histology in tadpoles appears to correlate with the TH activity, and in turn with the diversity in anuran life history transitions., (© 2019 Wiley Periodicals, Inc.)

The lateral line system in anurans is functional during aquatic stages and therefore could provide characters related to larval morphological variation. However, few studies have addressed its components in an integrated overview, and little is known about its ontogenetic variation. This study describes the postembryonic trajectory of the lateral system in Telmatobius atacamensis up to its metamorphic regression. This includes structure, number, topography, and innervation of neuromasts, to contribute new and complete information about its larval organization and its temporal sequence of regression. The arrangement and innervation of lateral lines in T. atacamensis resembles those described for other Type IV tadpoles. Its distinctive features are the orientation of the neuromast stitches in the lateral lines, the presence of supraotic neuromasts, and the first-described case of asymmetry of the ventral trunk line. The temporal sequence of regression during metamorphosis differs between the lateral lines and the lateral line nerves, which remain myelinated into postmetamorphic stages. This asynchronous pattern between different components of the system has also been described for Pseudis paradoxa, which shares with T. atacamensis a remarkably long larval period. This long larval period and gradual metamorphosis could also be related to the constitutive metamorphic regression of the system, in spite of the aquatic lifestyle of these frogs., (© 2019 Wiley Periodicals, Inc.)

Skeletochronology is the most accurate method to estimate a population age structure. The methodology is based on the analysis of secondary bone tissue in order to count growth rings. This study aimed to provide initial data, allowing researchers to further work out in the age of individuals and populations, sampling evidence of the presence of growth rings in 11 native species (representing nine families) of a subtropical region of southern Brazil. Four bone samples of each specimen were used to perform the skeletochronological analysis: the penultimate phalanges of the 3rd and 4th fingers, the humerus, and the femur. The presence of growth rings was confirmed in the periosteal layer of the bones of all analyzed species. In comparison with phalanges, growth rings of humeri and femora are more irregular and less distinguishable. This is the first record of growth rings to the native species herein analised. The skeletochronology was proved to be an effective tool in determining the age of anuran amphibians from a subtropical region, since this environment presents well defined climatic seasonality.

Genome size varies widely among organisms and is known to affect vertebrate development, morphology, and physiology. In amphibians, genome size is hypothesized to contribute to loss of late-forming structures, although this hypothesis has mainly been discussed in salamanders. Here we estimated genome size for 22 anuran species and combined this novel data set with existing genome size data for an additional 234 anuran species to determine whether larger genome size is associated with loss of a late-forming anuran sensory structure, the tympanic middle ear. We established that genome size is negatively correlated with development rate across 90 anuran species and found that genome size evolution is correlated with evolutionary loss of the middle ear bone (columella) among 241 species (224 eared and 17 earless). We further tested whether the development of the tympanic middle ear could be constrained by large cell sizes and small body sizes during key stages of tympanic middle ear development (metamorphosis). Together, our evidence suggests that larger genomes, slower development rate, and smaller body sizes at metamorphosis may contribute to the loss of the anuran tympanic middle ear. We conclude that increases in anuran genome size, although less drastic than those in salamanders, may affect development of late-forming traits.

Tree frogs are an interesting and diverse group of frogs. They display a number of unique adaptations to life in the arboreal environment. Vision plays a crucial role in their ecology. The topography of retinal ganglion cells (GCs) is closely related to a species' visual behavior. Despite a large amount of research addressing GC topography in vertebrates, there is scarce data on this subject in tree frogs. I studied the topography of GCs in the retina of the Japanese tree frog Hyla japonica. The GC density distribution was locally fairly homogeneous, with spatial density increasing gradually from the dorsal and ventral periphery towards the equator. A moderately pronounced visual streak was found close to the equator in the dorsal hemiretina, with a distinct area retinae temporalis in the dorsotemporal quadrant potentially subserving binocular vision. The minimum GC density (mean ± SEM, n = 5) was 3060 ± 60 and the maximum 12 800 ± 170 cells/mm 2 . The total number of GCs was 292 ± 7 × 10 3 . The theoretical anatomical spatial resolution estimated from GC densities and eye optics was lowest in the ventral periphery (ca. 0.9 and 1.3 cycles/degree in air and water, respectively) and highest in the area retinae temporalis (ca. 2.1 and 2.8 cycles/degree). The relatively high GC density and presence of specialized retinal regions in Hyla japonica are consistent with its highly visual behavior. The present findings contribute to our understanding of the relative role of common ancestry and environmental pressure in GC topography variation within Anura., (© 2019 Anatomical Society.)