Estratégias e modos reprodutivos de anuros (Amphibia) em uma poça permanente na Serra de Paranapiacaba, Sudeste do Brasil

Strategies and reproductive modes of anurans (Amphibia) in a permanent pond in Serra de Paranapiacaba, southeastern Brazil. This study describes the temporal patterns, behavioral strategies, reproductive modes, and fecundity of an anuran assemblage in a permanent pond in Serra de Paranapiacaba, municipality of Ribeirão Branco, south of São Paulo State (approximately 24º13'S; 48º46'W; ca. 800m above sea level). Field work was carried out between January and December 1993, totaling 40 nights of observation. Seven preliminary visits were made before this period and seven additional ones after December 1993. Naturalistic observations usually began before sunset and were concluded around 24:0001:00h. On three occasions the observations lasted the entire night. The commonest reproductive strategy among males of different species was that performed by calling male, but the satellite behavior was observed in some species. The calling temporal pattern was prolonged for the species of the assemblage; however, some species called only occasionally. Six different reproductive modes were observed in the pond or on its margins. The SVL of females of different species was positively correlated with egg clutch volume, irrespective of the reproductive mode. The SVL of females was also positively correlated with the number of eggs per clutch, although the correlation is stronger for species with a generalized reproductive mode. However, for species with specialized reproductive mode the number of eggs per clutch was not correlated with the SVL of females. A significant and positive correlation between female SVL and egg diameter was also detected for species with generalized reproductive mode. On the other hand, for species with specialized reproductive mode, female's SVL and egg diameter were negatively correlated. Terrestrial reproductive modes are possibly restricted to small-sized species. Large eggs may face problems of gas exchange due to their longer development period and lower surface/volume ratio.Este estudo descreve o padrão temporal, as estratégias reprodutivas, os modos reprodutivos e a fecundidade de uma taxocenose de anuros em uma poça permanente na Serra de Paranapiacaba, município de Ribeirão Branco, sul do estado de São Paulo (aproximadamente 24º13'S; 48º46'W; ca. 800m do nível do mar). O Trabalho de campo foi conduzido entre janeiro e dezembro de 1993, totalizando 40 noites de observações. Sete visitas preliminares foram feitas antes deste período e sete após. Observações naturalísticas normalmente se iniciavam antes do pôr do sol e eram concluídas aproximadamente entre 24:0001:00h. Em três ocasiões as observações foram ao longo da noite. A estratégia reprodutiva mais comum entre os machos de diferentes espécies foi a do macho cantor, mas comportamento satélite foi observado em algumas espécies. O padrão temporal foi prolongado para as espécies desta taxocenose; entretanto, algumas espécies podiam vocalizar ocasionalmente. Seis diferentes modos reprodutivos foram observados na poça ou suas margens. O comprimento rostro-cloacal (CRC) das fêmeas de diferentes espécies foi positivamente correlacionado com o volume da desova, independente do modo reprodutivo. O CRC das fêmeas foi positivamente correlacionado com o número de ovos por desova, embora a correlação seja mais forte para espécies com um modo reprodutivo generalizado. Entretanto, para espécies com modo reprodutivo especializado o número de ovos por desova não foi correlacionado com o CRC das fêmeas. Uma correlação positiva e significativa entre o CRC das fêmeas e o diâmetro dos ovos, também foi detectada para espécies com modo reprodutivo generalizado. Por outro lado, para as espécies com modo reprodutivo especializado o CRC das fêmeas e o diâmetro dos ovos foram negativamente correlacionados. Modos reprodutivos terrestres são possivelmente restritos a espécies pequenas. Ovos grandes podem apresentar problemas com a troca de gases devido ao seu longo desenvolvimento e baixa relação superfície/volume

Repertório vocal de Hylodes phyllodes (Amphibia, Anura, Hylodidae)

Hyllodes phyllodes is a diurnal frog, found in mountain streams in the Atlantic forest. Five vocalization types of H. phyllodes are described here, along with the social behavior with which they are associated. These five vocalizations include two types of advertisement calls, a courtship call, a territorial call, and a call emitted during the intervals between male-male conflicts. Frogs were studied from January 2001 to November 2002, in riverine creeks of the Atlantic forest, at Núcleo Picinguaba, Parque Estadual da Serra do Mar, Municipality of Ubatuba, state of São Paulo. Frogs were observed throughout the day, for a total of 80 hours of observations and recordings of 22 individuals. The advertisement call in this study is slightly different than that originally described, and may be due to environmental conditions, or behavioral variations, at the time of recording. Vocal adaptations of diurnal species that live in a noisy aquatic environment permit aural communication in a range of sounds outside those of the water noises. Thus, the calls of H. phyllodes have a dominant frequency much higher than that of the noise of the water current - that is, greater than 3.5 kHz. We show here that H. phyllodes has the richest known vocal repertoire within the genus Hylodes.Hyllodes phyllodes é um anfíbio diurno, que ocorre em córregos de montanha associados à Floresta Atlântica. Neste estudo, são caracterizados cinco tipos de vocalizações de Hylodes phyllodes relacionadas com o comportamento social desta espécie: dois tipos de canto de anúncio, canto de corte, canto territorial e canto de intervalo de briga. O estudo foi realizado entre janeiro de 2001 e novembro de 2002 em córregos na Mata Atlântica, dentro da área do Núcleo Picinguaba, Parque Estadual da Serra do Mar, no Município de Ubatuba, Estado de São Paulo. As observações foram realizadas em vários horários do dia, totalizando 80 horas de observações e gravação de 22 espécimes. O canto de anúncio descrito neste estudo apresentou pequenas diferenças em relação ao canto descrito originalmente, mas estas diferenças provavelmente são resultado de condições ambientais e/ou características comportamentais dos indivíduos gravados em Picinguaba e Boracéia. Adaptações na vocalização de espécies diurnas que vivem em riachos de correnteza permitem a comunicação sonora em um canal livre do barulho d'água, deste modo, os cantos de H. phyllodes têm freqüência dominante acima do barulho da correnteza, ou seja, maior que 3,5 kHz. Este estudo identifica H. phyllodes como a espécie com o repertório vocal mais rico do gênero

Tonal calls as a bioacoustic novelty in two Atlantic Forest species of Physalaemus (Anura: Leptodactylidae)

The frog genus Physalaemus has almost 50 species with vocalizations that are mostly composed of a single note. This note tends to have a broad harmonic structure or a pulsed structure. The sister species P. lateristriga and P. olfersii have pulsed advertisement calls that have been described as a noisy and long-lasting warbling sound. We provide the first account of inclusion of tonal sounds as part of the vocal repertoire of these species. Pure tones can (1) be long and form the entire call; (2) form prefixes of variable length separated by silence from the advertisement call; (3) be brief and form the onset or the offset of the regular advertisement call. Tonal calls may be an evolutionary novelty and they are not known from other populations of P. olfersii and P. lateristriga. Identification of the mechanism of sound production and of the behavioural role of these unique calls may help elucidate the evolution of call complexity in frogs

An extraordinary new species of melanophryniscus (anura, bufonidae) from Southeastern Brazil

We describe a new species of bufonid from a lowland, sandy soil, restinga habitat in the state of Espírito Santo, southeastern Brazil. Based on the shared occurrence of putative morphological synapomorphies of Melanophryniscus and the results of a phylogenetic analysis of DNA sequences of a broad sample of bufonids, and other anurans, we assign the new species to Melanophryniscus. The new species possesses several peculiar character states that distinguish it from all other Melanophryniscus including, but not limited to: fingers II, III, and V much reduced; nuptial pad with few enlarged, brown-colored spines on medial margin of finger II; seven presacral vertebrae, the last fused with the sacrum; and ventral humeral crest prominent, forming a spinelike projection. © Copyright © American Museum of Natural History 2012.Fil: Peloso, Pedro L.V.. American Museum of Natural History; Estados UnidosFil: Faivovich, Julián. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. American Museum of Natural History; Estados UnidosFil: Grant, Taran. American Museum of Natural History; Estados UnidosFil: Gasparini, João Luiz. Universidade Federal do Espírito Santo; BrasilFil: Haddad, Célio F.B.. Universidade Estadual Paulista Julio de Mesquita Filho; Brasi

A new genus of Cophomantini, with comments on the taxonomic status of Boana liliae (Anura: Hylidae)

© 2018 The Linnean Society of London The non-monophyly of both the genus Myersiohyla and the Boana punctata group has been recovered in a number of published phylogenetic analyses. In this paper we report on the analysis of sequences of Boana liliae, a species originally assigned to the B. punctata group, in a dataset of Cophomantini that recovered novel phylogenetic relationships for this hylid tribe. Our results reveal Myersiohyla to be paraphyletic with respect to B. liliae. Support for the placement of Myersiohyla kanaima is poor, but this taxon is recovered as the sister taxon of the other Cophomantini genera (excluding Myersiohyla) or as the sister taxon of the remaining species of Myersiohyla (including B. liliae). These results lead us to propose two taxonomic changes in order to remedy the paraphyly of Myersiohyla: (1) a new genus is described for M. kanaima, and (2) Boana liliae is transferred to Myersiohyla. We further provide notes on the natural history and vocalizations of the new monotypic genus, a new diagnosis of the former B. liliae in the context of Myersiohyla, and discuss the evolution of tadpole morphology and biogeography of the earlier diverging clades of Cophomantini

THE AMPHIBIAN TREE OF LIFE

The evidentiary basis of the currently accepted classification of living amphibians is discussed and shown not to warrant the degree of authority conferred on it by use and tradition. A new taxonomy of living amphibians is proposed to correct the deficiencies of the old one. This new taxonomy is based on the largest phylogenetic analysis of living Amphibia so far accomplished. We combined the comparative anatomical character evidence of Haas (2003) with DNA sequences from the mitochondrial transcription unit H1 (12S and 16S ribosomal RNA and tRNAValine genes, ø 2,400 bp of mitochondrial sequences) and the nuclear genes histone H3, rhodopsin, tyrosinase, and seven in absentia, and the large ribosomal subunit 28S (ø 2,300 bp of nuclear sequences; ca. 1.8 million base pairs; x¯ 5 3.7 kb/terminal). The dataset includes 532 terminals sampled from 522 species representative of the global diversity of amphibians as well as seven of the closest living relatives of amphibians for outgroup comparisons. The primary purpose of our taxon sampling strategy was to provide strong tests of the monophyly of all ‘‘family-group’’ taxa. All currently recognized nominal families and subfamilies were sampled, with the exception of Protohynobiinae (Hynobiidae). Many of the currently recognized genera were also sampled. Although we discuss the monophyly of genera, and provide remedies for nonmonophyly where possible, we also make recommendations for future research. A parsimony analysis was performed under Direct Optimization, which simultaneously optimizes nucleotide homology (alignment) and tree costs, using the same set of assumptions throughout the analysis. Multiple search algorithms were run in the program POY over a period of seven months of computing time on the AMNH Parallel Computing Cluster. Results demonstrate that the following major taxonomic groups, as currently recognized, are nonmonophyletic: Ichthyophiidae (paraphyletic with respect to Uraeotyphlidae), Caeciliidae (paraphyletic with respect to Typhlonectidae and Scolecomorphidae), Salamandroidea (paraphyletic with respect to Sirenidae), Leiopelmatanura (paraphyletic with respect to Ascaphidae), Discoglossanura (paraphyletic with respect to Bombinatoridae), Mesobatrachia (paraphyletic with respect to Neobatrachia), Pipanura (paraphyletic with respect to Bombinatoridae and Discoglossidae/Alytidae), Hyloidea (in the sense of containing Heleophrynidae; paraphyletic with respect to Ranoidea), Leptodactylidae (polyphyletic, with Batrachophrynidae forming the sister taxon of Myobatrachidae 1 Limnodynastidae, and broadly paraphyletic with respect to Hemiphractinae, Rhinodermatidae, Hylidae, Allophrynidae, Centrolenidae, Brachycephalidae, Dendrobatidae, and Bufonidae), Microhylidae (polyphyletic, with Brevicipitinae being the sister taxon of Hemisotidae), Microhylinae (poly/paraphyletic with respect to the remaining non-brevicipitine microhylids), Hyperoliidae (para/polyphyletic, with Leptopelinae forming the sister taxon of Arthroleptidae 1 Astylosternidae), Astylosternidae (paraphyletic with respect to Arthroleptinae), Ranidae (paraphyletic with respect to Rhacophoridae and Mantellidae). In addition, many subsidiary taxa are demonstrated to be nonmonophyletic, such as (1) Eleutherodactylus with respect to Brachycephalus; (2) Rana (sensu Dubois, 1992), which is polyphyletic, with various elements falling far from each other on the tree; and (3) Bufo, with respect to several nominal bufonid genera. A new taxonomy of living amphibians is proposed, and the evidence for this is presented to promote further investigation and data acquisition bearing on the evolutionary history of amphibians. The taxonomy provided is consistent with the International Code of Zoological Nomenclature (ICZN, 1999). Salient features of the new taxonomy are (1) the three major groups of living amphibians, caecilians/Gymnophiona, salamanders/Caudata, and frogs/Anura, form a monophyletic group, to which we restrict the name Amphibia; (2) Gymnophiona forms the sister taxon of Batrachia (salamanders 1 frogs) and is composed of two groups, Rhinatrematidae and Stegokrotaphia; (3) Stegokrotaphia is composed of two families, Ichthyophiidae (including Uraeotyphlidae) and Caeciliidae (including Scolecomorphidae and Typhlonectidae, which are regarded as subfamilies); (4) Batrachia is a highly corroborated monophyletic group, composed of two taxa, Caudata (salamanders) and Anura (frogs); (5) Caudata is composed of two taxa, Cryptobranchoidei (Cryptobranchidae and Hynobiidae) and Diadectosalamandroidei new taxon (all other salamanders); (6) Diadectosalamandroidei is composed of two taxa, Hydatinosalamandroidei new taxon (composed of Perennibranchia and Treptobranchia new taxon) and Plethosalamandroidei new taxon; (7) Perennibranchia is composed of Proteidae and Sirenidae; (8) Treptobranchia new taxon is composed of two taxa, Ambystomatidae (including Dicamptodontidae) and Salamandridae; (9) Plethosalamandroidei new taxon is composed of Rhyacotritonidae and Xenosalamandroidei new taxon; (10) Xenosalamandroidei is composed of Plethodontidae and Amphiumidae; (11) Anura is monophyletic and composed of two clades, Leiopelmatidae (including Ascaphidae) and Lalagobatrachia new taxon (all other frogs); (12) Lalagobatrachia is composed of two clades, Xenoanura (Pipidae and Rhinophrynidae) and Sokolanura new taxon (all other lalagobatrachians); (13) Bombinatoridae and Alytidae (former Discoglossidae) are each others’ closest relatives and in a clade called Costata, which, excluding Leiopelmatidae and Xenoanura, forms the sister taxon of all other frogs, Acosmanura; (14) Acosmanura is composed of two clades, Anomocoela (5 Pelobatoidea of other authors) and Neobatrachia; (15) Anomocoela contains Pelobatoidea (Pelobatidae and Megophryidae) and Pelodytoidea (Pelodytidae and Scaphiopodidae), and forms the sister taxon of Neobatrachia, together forming Acosmanura; (16) Neobatrachia is composed of two clades, Heleophrynidae, and all other neobatrachians, Phthanobatrachia new taxon; (17) Phthanobatrachia is composed of two major units, Hyloides and Ranoides; (18) Hyloides comprises Sooglossidae (including Nasikabatrachidae) and Notogaeanura new taxon (the remaining hyloids); (19) Notogaeanura contains two taxa, Australobatrachia new taxon and Nobleobatrachia new taxon; (20) Australobatrachia is a clade composed of Batrachophrynidae and its sister taxon, Myobatrachoidea (Myobatrachidae and Limnodynastidae), which forms the sister taxon of all other hyloids, excluding sooglossids; (21) Nobleobatrachia new taxon, is dominated at its base by frogs of a treefrog morphotype, several with intercalary phalangeal cartilages—Hemiphractus (Hemiphractidae) forms the sister taxon of the remaining members of this group, here termed Meridianura new taxon; (22) Meridianura comprises Brachycephalidae (former Eleutherodactylinae 1 Brachycephalus) and Cladophrynia new taxon; (23) Cladophrynia is composed of two groups, Cryptobatrachidae (composed of Cryptobatrachus and Stefania, previously a fragment of the polyphyletic Hemiphractinae) and Tinctanura new taxon; (24) Tinctanura is composed of Amphignathodontidae (Gastrotheca and Flectonotus, another fragment of the polyphyletic Hemiphractinae) and Athesphatanura new taxon; (25) Athesphatanura is composed of Hylidae (Hylinae, Pelodryadinae, and Phyllomedusinae, and excluding former Hemiphractinae, whose inclusion would have rendered this taxon polyphyletic) and Leptodactyliformes new taxon; (26) Leptodactyliformes is composed of Diphyabatrachia new taxon (composed of Centrolenidae [including Allophryne] and Leptodactylidae, sensu stricto, including Leptodactylus and relatives) and Chthonobatrachia new taxon; (27) Chthonobatrachia is composed of a reformulated Ceratophryidae (which excludes such genera as Odontophrynus and Proceratophrys and includes other taxa, such as Telmatobius) and Hesticobatrachia new taxon; (28) Hesticobatrachia is composed of a reformulated Cycloramphidae (which includes Rhinoderma) and Agastorophrynia new taxon; (29) Agastorophrynia is composed of Bufonidae (which is partially revised) and Dendrobatoidea (Dendrobatidae and Thoropidae); (30) Ranoides new taxon forms the sister taxon of Hyloides and is composed of two major monophyletic components, Allodapanura new taxon (microhylids, hyperoliids, and allies) and Natatanura new taxon (ranids and allies); (31) Allodapanura is composed of Microhylidae (which is partially revised) and Afrobatrachia new taxon; (32) Afrobatrachia is composed of Xenosyneunitanura new taxon (the ‘‘strange-bedfellows’’ Brevicipitidae [formerly in Microhylidae] and Hemisotidae) and a more normal-looking group of frogs, Laurentobatrachia new taxon (Hyperoliidae and Arthroleptidae, which includes Leptopelinae and former Astylosternidae); (33) Natatanura new taxon is composed of two taxa, the African Ptychadenidae and the worldwide Victoranura new taxon; (34) Victoranura is composed of Ceratobatrachidae and Telmatobatrachia new taxon; (35) Telmatobatrachia is composed of Micrixalidae and a worldwide group of ranoids, Ametrobatrachia new taxon; (36) Ametrobatrachia is composed of Africanura new taxon and Saukrobatrachia new taxon; (37) Africanura is composed of two taxa: Phrynobatrachidae (Phrynobatrachus, including Dimorphognathus and Phrynodon as synonyms) and Pyxicephaloidea; (38) Pyxicephaloidea is composed of Petropedetidae (Conraua, Indirana, Arthroleptides, and Petropedetes), and Pyxicephalidae (including a number of African genera, e.g. Amietia [including Afrana], Arthroleptella, Pyxicephalus, Strongylopus, and Tomopterna); and (39) Saukrobatrachia new taxon is the sister taxon of Africanura and is composed of Dicroglossidae and Aglaioanura new taxon, which is, in turn, composed of Rhacophoroidea (Mantellidae and Rhacophoridae) and Ranoidea (Nyctibatrachidae and Ranidae, sensu stricto). Many generic revisions are made either to render a monophyletic taxonomy or to render a taxonomy that illuminates the problems in our understanding of phylogeny, so that future work will be made easier. These revisions are: (1) placement of Ixalotriton and Lineatriton (Caudata: Plethodontidae: Bolitoglossinae) into the synonymy of Pseudoeurycea, to render a monophyletic Pseudoeurycea; (2) placement of Haideotriton (Caudata: Plethodontidae: Spelerpinae) into the synonymy of Eurycea, to render a monophyletic Eurycea; (3) placement of Nesomantis (Anura: Sooglossidae) into the synonymy of Sooglossus, to assure a monophyletic Sooglossus; (4) placement of Cyclorana and Nyctimystes (Anura: Hylidae: Pelodryadinae) into Litoria, but retaining Cyclorana as a subgenus, to provide a monophyletic Litoria; (5) partition of ‘‘Limnodynastes’’ (Anura: Limnodynastidae) into Limnodynastes and Opisthodon to render monophyletic genera; (6) placement of Adenomera, Lithodytes, and Vanzolinius (Anura: Leptodactylidae) into Leptodactylus, to render a monophyletic Leptodactylus; (7) partition of ‘‘Eleutherodactylus’’ (Anura: Brachycephalidae) into Craugastor, ‘‘Eleutherodactylus’’, ‘‘Euhyas’’, ‘‘Pelorius’’, and Syrrhophus to outline the taxonomic issues relevant to the paraphyly of this nominal taxon to other nominal genera; (8) partition of ‘‘Bufo’’ (Anura: Bufonidae) into a number of new or revived genera (i.e., Amietophrynus new genus, Anaxyrus, Chaunus, Cranopsis, Duttaphrynus new genus, Epidalea, Ingerophrynus new genus, Nannophryne, Peltophryne, Phrynoidis, Poyntonophrynus new genus; Pseudepidalea new genus, Rhaebo, Rhinella, Vandijkophrynus new genus); (9) placement of the monotypic Spinophrynoides (Anura: Bufonidae) into the synonymy of (formerly monotypic) Altiphrynoides to make for a more informative taxonomy; (10) placement of the Bufo taitanus group and Stephopaedes (as a subgenus) into the synonymy of Mertensophryne (Anura: Bufonidae); (11) placement of Xenobatrachus (Anura: Microhylidae: Asterophryinae) into the synonymy of Xenorhina to render a monophyletic Xenorhina; (12) transfer of a number of species from Plethodontohyla to Rhombophryne (Microhylidae: Cophylinae) to render a monophyletic Plethodontohyla; (13) placement of Schoutedenella (Anura: Arthroleptidae) into the synonymy of Arthroleptis; (14) transfer of Dimorphognathus and Phrynodon (Anura: Phrynobatrachidae) into the synonymy of Phrynobatrachus to render a monophyletic Phrynobatrachus; (15) placement of Afrana into the synonymy of Amietia (Anura: Pyxicephalidae) to render a monophyletic taxon; (16) placement of Chaparana and Paa into the synonymy of Nanorana (Anura: Dicroglossidae) to render a monophyletic genus; (17) recognition as genera of Ombrana and Annandia (Anura: Dicroglossidae: Dicroglossinae) pending placement of them phylogenetically; (18) return of Phrynoglossus into the synonymy of Occidozyga to resolve the paraphyly of Phrynoglossus (Anura: Dicroglossidae: Occidozyginae); (19) recognition of Feihyla new genus for Philautus palpebralis to resolve the polyphyly of ‘‘Chirixalus’’; (20) synonymy of ‘‘Chirixalus’’ with Chiromantis to resolve the paraphyly of ‘‘Chirixalus’’; (21) recognition of the genus Babina, composed of the former subgenera of Rana, Babina and Nidirana (Anura: Ranidae); (22) recognition of the genera Clinotarsus, Humerana, Nasirana, Pelophylax, Pterorana, Pulchrana, and Sanguirana, formerly considered subgenera of Rana (Anura: Ranidae), with no special relationship to Rana (sensu stricto); (23) consideration of Glandirana (Anura: Ranidae), formerly a subgenus of Rana, as a genus, with Rugosa as a synonym; (24) recognition of Hydrophylax (Anura: Ranidae) as a genus, with Amnirana and most species of former Chalcorana included in this taxon as synonyms; (25) recognition of Hylarana (Anura: Ranidae) as a genus and its content redefined; (26) redelimitation of Huia to include as synonyms Eburana and Odorrana (both former subgenera of Rana); (27) recognition of Lithobates (Anura: Ranidae) for all species of North American ‘‘Rana’’ not placed in Rana sensu stricto (Aquarana, Pantherana, Sierrana, Trypheropsis, and Zweifelia considered synonyms of Lithobates); (28) redelimitation of the genus Rana as monophyletic by inclusion as synonyms Amerana, Aurorana, Pseudoamolops, and Pseudorana, and exclusion of all other former subgenera; (29) redelimitation of the genus Sylvirana (Anura: Ranidae), formerly a subgenus of Rana, with Papurana and Tylerana included as synonyms

Multiple connections between Amazonia and Atlantic Forest shaped the T phylogenetic and morphological diversity of Chiasmocleis Mehely, 1904 (Anura: Microhylidae: Gastrophryninae)

Chiasmocleis is the most species-rich genus of Neotropical microhylids. Herein, we provide the first comprehensive multilocus phylogeny for the genus, including all but 3 of the 34 recognized species and multiple individuals per species. We discuss cryptic speciation, species discovery, patterns of morphological evolution, and provide a historical biogeographic analysis to account for the current distribution of the genus. Diversification of Chiasmocleis from other New World microhylids began during the Eocene, app. 40 mya, in forested areas, and current diversity seems to be a product of recurrent connections between the Atlantic Forest and Amazonia. Small-sized species evolved independently three times in Chiasmocleis. Furthermore, the extremely small-bodied (i.e. miniaturized) species with associated loss of digits, phalanges, and pectoral girdle cartilages evolved only once and are restricted to Amazonia. Using the phylogeny, we recognized three subgenera within Chiasmocleis: Chiasmocleis Méhely, 1904, Relictus subg. nov., and Syncope Walker, 1973. The recognition of the subgenus Syncopeinforms future research on patterns of miniaturization in the genus, and the subgenus Relictus highlights isolation of an endemic and species-poor lineage to the Atlantic Forest, early (about 40 mya) in the history of Chiasmocleis

Evolution in the Genus Rhinella: A Total Evidence Phylogenetic Analysis of Neotropical True Toads (Anura: Bufonidae)

True toads of the genus Rhinella are among the most common and diverse group of Neotropical anurans. These toads are widely distributed throughout South America, inhabiting a great diversity of environments and ecoregions. Currently, however, the genus is defined solely on the basis of molecular characters, and it lacks a proper diagnosis. Although some phenetic species groups have traditionally been recognized within Rhinella, the monophyly of some of them have been rejected in previous phylogenetic analyses, and many species remain unassigned to these poorly defined groups. Additionally, the identity and taxonomy of several species are problematic and hinder the specific recognition and description of undescribed taxa. In this work, we first perform phylogenetic analyses of separate mitochondrial and nuclear datasets to test the possible occurrence of hybridiza-tion and/or genetic introgression in the genus. The comparative analysis of both datasets revealed unidirectional mitochondrial introgressions of an unknown parental species into R . horribilis (“ghost introgression”) and of R . dorbignyi into R . bernardoi; therefore, the mitochondrial and nuclear data-sets of these species were considered separately in subsequent analyses. We performed total-evidence phylogenetic analyses that included revised molecular (four mitochondrial and five nuclear genes) and phenotypic (90 characters) datasets for 83 nominal species of Rhinella, plus several undescribed and problematic species and multiple outgroups. Results demonstrate that Rhinella was nonmono-phyletic due to the position of R . ceratophrys, which was recovered as the sister taxon of Rhaebo nasicus with strong support. Among our outgroups, the strongly supported Anaxyrus + Incilius is the sister clade of all other species of Rhinella. Once R . ceratophrys is excluded, the genus Rhinellais monophyletic, well supported, and composed of two major clades. One of these is moderately supported and includes species of the former R . spinulosa Group (including R . gallardoi); the mono-phyletic R . granulosa, R . crucifer, and R . marina Groups; and a clade composed of the mitochondrial sequences of R . horribilis. The other major clade is strongly supported and composed of all the spe-cies from the non-monophyletic R . veraguensis and R . margaritifera Groups, the former R . acrolophaGroup, and R . sternosignata. Consistent with these results, we define eight species groups of Rhinella that are mostly diagnosed by phenotypic synapomorphies in addition to a combination of morpho-logical character states. Rhinella sternosignata is the only species that remains unassigned to any group. We also synonymize nine species, treat three former subspecies as full species, and suggest that 15 lineages represent putative undescribed species. Lastly, we discuss the apparently frequent occurrence of hybridization, deep mitochondrial divergence, and “ghost introgression”; the incomplete phenotypic evidence (including putative character systems that could be used for future phy-logenetic analyses); and the validity of the known fossil record of Rhinella as a source of calibration points for divergence dating analyses.Peer reviewe