Environment rather than character displacement explains call evolution in glassfrogs.

The acoustic adaptation hypothesis (AAH) and ecological character displacement (ECD) are two potential mechanisms shaping call evolution that can predict opposite trends for the differentiation of signals. Under AAH, signals evolve to minimize environmental degradation and maximize detection against background noise, predicting call homogenization in similar habitats due to environmental constraints on signals. In contrast, ECD predicts greater differences in call traits of closely related taxa in sympatry because of selection against acoustic interference. We used comparative phylogenetic analyses to test the strength of these two selective mechanisms on the evolution of advertisement calls in glassfrogs, a highly diverse family of neotropical anurans. We found that, overall, acoustic adaptation to the environment may outweigh effects of species interactions. As expected under the AAH, temporal call parameters are correlated with vegetation density, but spectral call parameters had an unexpected inverse correlation with vegetation density, as well as an unexpected correlation with temperature. We detected call convergence among co-occurring species and also across multiple populations from the same species in different glassfrogs communities. Our results indicate that call convergence is common in glassfrogs, likely due to habitat filtering, while character displacement is relatively rare, suggesting that costs of signal similarity among related species may not drive divergent selection in all systems.

Auditory Survey of Endangered Eurasian Bittern Using Microphone Arrays and Robot Audition.

Bioacoustics monitoring has become increasingly popular for studying the behavior and ecology of vocalizing birds. This study aims to verify the practical effectiveness of localization technology for auditory monitoring of endangered Eurasian bittern (Botaurus stellaris) which inhabits wetlands in remote areas with thick vegetation. Their crepuscular and highly secretive nature, except during the breeding season when they vocalize advertisement calls, make them difficult to monitor. Because of the increasing rates of habitat loss, surveying accurate numbers and their habitat needs are both important conservation tasks. We investigated the feasibility of localizing their booming calls, at a low frequency range between 100-200 Hz, using microphone arrays and robot audition HARK (Honda Research Institute, Audition for Robots with Kyoto University). We first simulated sound source localization of actual bittern calls for microphone arrays of radii 10 cm, 50 cm, 1 m, and 10 m, under different noise levels. Second, we monitored bitterns in an actual field environment using small microphone arrays (height = 12 cm; width = 8 cm), in the Sarobetsu Mire, Hokkaido Island, Japan. The simulation results showed that the spectral detectability was higher for larger microphone arrays, whereas the temporal detectability was higher for smaller microphone arrays. We identified that false detection in smaller microphone arrays, which was coincidentally generated in the calculation proximate to the transfer function for the opposite side. Despite technical limitations, we successfully localized booming calls of at least two males in a reverberant wetland, surrounded by thick vegetation and riparian trees. This study is the first case of localizing such rare birds using small-sized microphone arrays in the field, thereby presenting how this technology could contribute to auditory surveys of population numbers, behaviors, and microhabitat selection, all of which are difficult to investigate using other observation methods. This methodology is not only useful for the better understanding of bitterns, but it can also be extended to investigate other rare nocturnal birds with low-frequency vocalizations, without direct ringing or tagging. Our results also suggest a future necessity for a robust localization system to avoid reverberation and echoing in the field, resulting in the false detection of the target birds.

Conspecific odor cues induce different vocal responses in serrate-legged small treefrogs, but only in the absence of acoustic signals.

BACKGROUND: Signal detection is crucial to survival and successful reproduction, and animals often modify behavioral decisions based on information they obtained from the social context. Undeniably, the decision-making in male-male competition and female choice of anurans (frogs and toads) depends heavily on acoustic signals. However, increasing empirical evidence suggests that additional or alternative types of cue (e.g., visual, chemical, and vibratory) can be used to detect, discriminate and locate conspecifics in many anuran species. Nevertheless, few studies have investigated whether conspecific odor cues affect male's calling behavior. In this study, we conducted an experiment to investigate whether and how different chemical cues (male odors, female odors, and stress odors) from conspecifics affect male's calling strategies in serrate-legged small treefrogs (Kurixalus odontotarsus), and whether the combined chemical and acoustic stimuli have additive effects on calling behavior or not. RESULTS: We found that compared with female odors, male K. odontotarsus reduced calling investment in response to male odors or stress odors, in the absence of rival's advertisement calls. When odor stimuli and advertisement calls were presented simultaneously, however, there were no differences in the vocal response of focal males among odor groups. CONCLUSIONS: These results provide evidence that male treefrogs switch calling investment according to different odor cues from conspecifics, and further demonstrate that calling behavior can be affected by chemical cues in anuran species. Our study highlights the potential role of airborne chemical cues in sex identification and contributes to increase our understanding of anuran communication.

Vocalization by extant nonavian reptiles: A synthetic overview of phonation and the vocal apparatus.

Among amniote vertebrates, nonavian reptiles (chelonians, crocodilians, and lepidosaurs) are regarded as using vocal signals rarely (compared to birds and mammals). In all three reptilian clades, however, certain taxa emit distress calls and advertisement calls using modifications of regions of the upper respiratory tract. There is no central tendency in either acoustic mechanisms or the structure of the vocal apparatus, and many taxa that vocalize emit only relatively simple sounds. Available evidence indicates multiple origins of true vocal abilities within these lineages. Reptiles thus provide opportunities for studying the early evolutionary stages of vocalization. The early literature on the diversity of form of the laryngotracheal apparatus of reptiles boded well for the study of form-function relationships, but this potential was not extensively explored. Emphasis shifted away from anatomy, however, and centered instead on acoustic analysis of the sounds that are produced. New investigative techniques have provided novel ways of studying the form-function aspects of the structures involved in phonation and have brought anatomical investigation to the forefront again. In this review we summarize what is known about hearing in reptiles in order to contextualize the vocal signals they generate and the sound-producing mechanisms responsible for them. The diversity of form of the sound producing apparatus and the increasing evidence that reptiles are more dependent upon vocalization as a communication medium than previously thought indicates that they have a significant role to play in the understanding of the evolution of vocalization in amniotes.

Correction to the type locality of Tomopterna ahli (Deckert, 1938) (Anura: Pyxicephalidae), with the designation of a neotype.

Deckert described a Tomopterna as Arthroleptella ahli from South Africa in 1938. We demonstrate that the type of Tomopterna ahli (Deckert, 1938) was actually collected in Namibia, and that this nomen is a senior synonym of Tomopterna damarensis Dawood Channing, 2002. We designate a neotype for Tomopterna ahli (Deckert, 1938).

Acoustic niche partitioning in an anuran community from the municipality of Floriano, Piauí, Brazil / Partição de nicho acústico em uma comunidade de anuros do município de Floriano, Piauí, Brasil

Abstract Vocalizations are an important trait for the identification of cryptic and/or closely related amphibian species. Different vocalizations also contribute to partitioning of the acoustic space by sympatric species. This study aimed to describe the advertisement calls of anurans in a pond of the municipality of Floriano, State of Piauí, Brazil, and infer the acoustic niche partitioning of amphibians. Euclidean distance was used in a cluster analysis approach to infer the acoustic similarities among species. Thirteen species were analysed: Boana raniceps, Dendropsophus nanus, D. rubicundulus , D. minutus, Leptodactylus fuscus, L. troglodytes, L. vastus, Pithecopus nordestinus , Physalaemus cuvieri, P. nattereri, Pleurodema diplolister, Proceratophrys cristiceps and Scinax ruber. From these, six showed more than 90% of acoustic overlap: P. nattereri , P. cuvieri, L. fuscus and L. vastus (Leptodactylidae); and, D. nanus and D. rubicundulus (Hylidae). Despite the acoustic similarities among these six species, the acoustic interference was reduced due to the small number of sympatric species and to distinct features on carrier frequency such as dominant frequency and the degree of modulation in the frequency. Environmental factors limit the periods and sites of reproduction respectively, which may maintain the low anuran diversity and consequently reduce acoustic overlap.

Resumo A vocalização é um componente importante na identificação de espécies crípticas e/ou proximamente relacionadas de anfíbios anuros. Diferenças na bioacústica também contribuem para o particionamento do espaço acústico entre espécies simpátricas. Este estudo descreve o canto de anúncio de anuros em uma lagoa temporária do município de Floriano, Estado do Piauí, Brasil, além de buscar inferir o particionamento do nicho acústico entre essas espécies. Para tanto, foi utilizada a distância Euclidiana para compor uma análise de conglomerados e avaliar a similaridade/dissimilaridade da bioacústica da anurofauna. Treze espécies foram amostradas: Boana raniceps, Dendropsophus nanus, D. rubicundulus , D. minutus, Leptodactylus fuscus, L. troglodytes, L. vastus, Pithecopus nordestinus , Physalaemus cuvieri, P. nattereri, Pleurodema diplolister, Proceratophrys cristiceps e Scinax ruber. Destas, seis apresentaram mais de 90% de sobreposição acústica: P. nattereri, P. cuvieri, L. fuscus e L. vastus (Leptodactylidae); e, D. nanus e D. rubicundulus (Hylidae). Apesar da similaridade acústica entre essas seis espécies, a interferência é reduzida devido ao pequeno número de espécies simpátricas e certas características distintas na frequência do canto, tais como: frequência dominante e sua modulação. Fatores ambientais específicos do semiárido limitam os períodos e sítios reprodutivos, o que mantém baixa diversidade de espécies e consequentemente reduz a sobreposição no nicho acústico.

Low Geographic and Subspecific Variation in the Loud Call of the Widespread and Phenotypically Cryptic Northern Lesser Galago (Galago senegalensis) Suggests Taxonomic Uniformity.

Like other nocturnal primates, many species of galago (Galagidae) are phenotypically cryptic, making their taxonomic status difficult to resolve. Recent taxonomic work has disentangled some of the confusion. This has resulted in an increase in the number of recognised galago species. The most widespread galago species, and indeed the most widespread nocturnal primate, is the northern lesser galago (Galago senegalensis) whose geographic range stretches >7,000 km across Africa. Based on morphology, 4 subspecies are currently recognised: G. s. senegalensis, G. s. braccatus, G. s. sotikae and G. s. dunni. We explore geographic and subspecific acoustic variation in G. senegalensis, testing three hypotheses: isolation by distance, genetic basis, and isolation by barrier. There is statistical support for isolation by distance for 2 of 4 call parameters (fundamental frequency and unit length). Geographic distance explains a moderate amount of the acoustic variation. Discriminant function analysis provides some degree of separation of geographic regions and subspecies, but the percentage of misdesignation is high. Despite having (putative) parapatric geographic ranges, the most pronounced acoustic differences are between G. s. senegalensis and G. s. dunni. The findings suggest that the Eastern Rift Valley and Niger River are significant barriers for G. senegalensis. The acoustic structures of the loud calls of 121 individuals from 28 widespread sites are not significantly different. Although this makes it unlikely that additional unrecognised species occur within G. senegalensis at the sites sampled, vast areas of the geographic range remain unsampled. We show that wide-ranging species do not necessarily exhibit large amounts of variation in their vocal repertoire. This pattern may also be present in nocturnal primates with smaller geographic ranges.

A name for the nurse-frog (Allobates, Aromobatidae) of Floresta Nacional de Carajás, Eastern Brazilian Amazonia.

We describe a new species of nurse-frog (Aromobatidae, Allobates) from Floresta Nacional de Carajás, southeastern Brazilian Amazonia. Allobates carajas sp. nov. is distinguished from similar congeneric species by the combination of the following characters: body-size range (snout-to-vent length 16.5-19.1 mm), dorsal color pattern with a dark brown hourglass-shaped mark, by the absence of a pale dorsolateral stripe and presence of a pale ventrolateral stripe, absence of a pale paracloacal mark, by overall bright yellow ventral colors of live male and female specimens, and by the posterior labium of tadpoles, with a single row of pyramidal papillae medially. The advertisement call of the new species is highly variable, with four possible temporal arrangements of notes (continuous emission of notes separated by regular silent intervals, continuous emission of notes separated by irregular silent intervals, emission of discrete note trills, and sporadic emission of single notes). Duration of notes range between 0.020-0.060 s, and the dominant frequency of notes range from 4.75 to 5.38 kHz. The new species is currently known only from forested habitats within Floresta Nacional de Carajás.

A new miniature Melanesian Forest Frog (Ceratobatrachidae: Cornufer) from New Britain Island, constituting the first record of the subgenus Batrachylodes from outside of the Solomon Archipelago.

We describe a new species of Cornufer, subgenus Batrachylodes, from high-elevation forests of New Britain Island in the Bismarck Archipelago of Eastern Melanesia. The new species, Cornufer exedrus sp. nov., is a biogeographically disjunct member of the Batrachylodes clade, representing the first record of the subgenus from outside of the Solomon Archipelago. The new species is a small terrestrial form from dense, closed-canopy forests above 1500 meters elevation in the Nakanai Mountains of eastern New Britain. It differs from its closest relatives, the other members of the subgenus Batrachylodes, on the basis of its minute body size, degree of digital disc expansion, reduced subdigital tuberculation, color pattern, and other traits related to its small size. We also provide a description of the new species' simple advertisement call. The diversity of ceratobatrachid frogs of the Bismarck Archipelago is most likely still underestimated despite several recent surveys. Our understanding of evolutionary trends and species boundaries in the subgenus Batrachylodes currently is hampered by lack of genetic samples and call recordings corresponding to voucher specimens of the endemic species of Bougainville Island.

A new species of nurse-frog (Aromobatidae, Allobates) from the Juami River basin, northwestern Brazilian Amazonia.

We describe a new species of nurse-frog (Aromobatidae, Allobates) from northwestern Brazilian Amazon. Allobates juami sp. nov. is distinguished from similar congeneric species by the combination of the following characters: body-size range (snout-to-vent length 17.5-18.5 mm), lack of dark pigments on ventral surfaces of male specimens, dorsal color pattern (predominantly solid dark brown, but conspicuously light brown over snout and urostyle regions), presence of conspicuous pale dorsolateral and ventrolateral stripes, and presence of a diffuse pale paracloacal mark. The advertisement call of the new species lasts 2.5-5.1 s, contains 60-73 short notes (trills), and is emitted at an average rate of 13 notes per second within trills. Duration of silent intervals between notes ranges between 0.020-0.050 s, and the peak frequency of notes ranges from 4.59 to 5.47 kHz. The new species is currently known only from the type locality at Estação Ecológica Juami-Japurá (1.96455° S, 67.93579° W; ~ 87 m a.s.l.).

Acoustic niche partitioning in an anuran community from the municipality of Floriano, Piauí, Brazil

Abstract Vocalizations are an important trait for the identification of cryptic and/or closely related amphibian species. Different vocalizations also contribute to partitioning of the acoustic space by sympatric species. This study aimed to describe the advertisement calls of anurans in a pond of the municipality of Floriano, State of Piauí, Brazil, and infer the acoustic niche partitioning of amphibians. Euclidean distance was used in a cluster analysis approach to infer the acoustic similarities among species. Thirteen species were analysed: Boana raniceps, Dendropsophus nanus, D. rubicundulus , D. minutus, Leptodactylus fuscus, L. troglodytes, L. vastus, Pithecopus nordestinus , Physalaemus cuvieri, P. nattereri, Pleurodema diplolister, Proceratophrys cristiceps and Scinax ruber. From these, six showed more than 90% of acoustic overlap: P. nattereri , P. cuvieri, L. fuscus and L. vastus (Leptodactylidae); and, D. nanus and D. rubicundulus (Hylidae). Despite the acoustic similarities among these six species, the acoustic interference was reduced due to the small number of sympatric species and to distinct features on carrier frequency such as dominant frequency and the degree of modulation in the frequency. Environmental factors limit the periods and sites of reproduction respectively, which may maintain the low anuran diversity and consequently reduce acoustic overlap.

Resumo A vocalização é um componente importante na identificação de espécies crípticas e/ou proximamente relacionadas de anfíbios anuros. Diferenças na bioacústica também contribuem para o particionamento do espaço acústico entre espécies simpátricas. Este estudo descreve o canto de anúncio de anuros em uma lagoa temporária do município de Floriano, Estado do Piauí, Brasil, além de buscar inferir o particionamento do nicho acústico entre essas espécies. Para tanto, foi utilizada a distância Euclidiana para compor uma análise de conglomerados e avaliar a similaridade/dissimilaridade da bioacústica da anurofauna. Treze espécies foram amostradas: Boana raniceps, Dendropsophus nanus, D. rubicundulus , D. minutus, Leptodactylus fuscus, L. troglodytes, L. vastus, Pithecopus nordestinus , Physalaemus cuvieri, P. nattereri, Pleurodema diplolister, Proceratophrys cristiceps e Scinax ruber. Destas, seis apresentaram mais de 90% de sobreposição acústica: P. nattereri, P. cuvieri, L. fuscus e L. vastus (Leptodactylidae); e, D. nanus e D. rubicundulus (Hylidae). Apesar da similaridade acústica entre essas seis espécies, a interferência é reduzida devido ao pequeno número de espécies simpátricas e certas características distintas na frequência do canto, tais como: frequência dominante e sua modulação. Fatores ambientais específicos do semiárido limitam os períodos e sítios reprodutivos, o que mantém baixa diversidade de espécies e consequentemente reduz a sobreposição no nicho acústico.

Geographic variation in the advertisement calls of Hyla eximia and its possible explanations.

Populations of species occupying large geographic ranges are often phenotypically diverse as a consequence of variation in selective pressures and drift. This applies to attributes involved in mate choice, particularly when both geographic range and breeding biology overlap between related species. This condition may lead to interference of mating signals, which would in turn promote reproductive character displacement (RCD). We investigated whether variation in the advertisement call of the mountain treefrog (Hyla eximia) is linked to geographic distribution with respect to major Mexican river basins (Panuco, Lerma, Balsas and Magdalena), or to coexistence with its sister (the canyon treefrog, Hyla arenicolor) or another related species (the dwarf treefrog, Tlalocohyla smithii). We also evaluated whether call divergence across the main river basins could be linked to genetic structure. We found that the multidimensional acoustic space of calls from two basins where H. eximia currently interacts with T. smithii, was different from the acoustic space of calls from H. eximia elsewhere. Individuals from these two basins were also distinguishable from the rest by both the phylogeny inferred from mitochondrial sequences, and the genetic structure inferred from nuclear markers. The discordant divergence of H. eximia advertisement calls in the two separate basins where its geographic range overlaps that of T. smithii can be interpreted as the result of two independent events of RCD, presumably as a consequence of acoustic interference in the breeding choruses, although more data are required to evaluate this possibility.

Acoustic characteristics of eight common Chinese anurans during the breeding season.

Anurans often have species-specific vocalizations. To quantify and compare the characteristics of anuran calls in Gutianshan National Nature Reserve, Zhejiang Province, we recorded the advertisement calls of eight species belonging to four families (Ranidae, Microhylidae, Megophryidae and Bufonidae) from June to September 2012 using Sony ICD-FX8 IC recorders. All recordings were analyzed using the "Praat" software. Five acoustics parameters were measured, including temporal traits (call duration, number of notes or pulse number/call) and spectral traits (fundamental frequency, the first three formants and dominant frequency). The characteristic parameters of Microhyla ornate and Fejervarya limnocharis calls were different as were the calls of some populations of the same species recorded in different regions. The advertisement calls of the eight species were specific. Our study has provided a useful reference for identifying the calls of some common Chinese anurans.

Three new species of Oreophryne (Anura, Microhylidae) from Papua New Guinea.

I describe three new species of the diverse microhylid frog genus Oreophryne from Papua New Guinea. Two of these occur in two isolated mountain ranges along the northern coast of Papua New Guinea; the third is from Rossel Island in the very southeasternmost part of the country. All three are the first Oreophryne known from these areas to have a cartilaginous connection between the procoracoid and scapula, a feature usually seen in species far to the west or from the central cordillera of New Guinea. Each of the new species also differs from the many other Papuan Oreophryne in a variety of other morphological, color-pattern, and call features. Advertisement-call data for Oreophryne species from the north-coast region suggest that they represent only two of the several call types seen in regions further south, consistent with the relatively recent derivation of these northern regions as accreted island-arc systems. The distinctively different, whinnying, call type of the new species from Rossel Island occurs among other Oreophryne from southeastern Papua New Guinea but has been unreported elsewhere, raising the possibility that it may characterize a clade endemic to that region.

Systematics of the Osteocephalusbuckleyi species complex (Anura, Hylidae) from Ecuador and Peru.

We present a new phylogeny, based on DNA sequences of mitochondrial and nuclear genes, for frogs of the genus Osteocephalus with emphasis in the Osteocephalus buckleyi species complex. Genetic, morphologic, and advertisement call data are combined to define species boundaries and describe new species. The phylogeny shows strong support for: (1) a basal position of Osteocephalus taurinus + Osteocephalus oophagus, (2) a clade containing phytotelmata breeding species, and (3) a clade that corresponds to the Osteocephalus buckleyi species complex. Our results document a large proportion of hidden diversity within a set of populations that were previously treated as a single, widely distributed species, Osteocephalus buckleyi. Individuals assignable to Osteocephalus buckleyi formed a paraphyletic group relative to Osteocephalus verruciger and Osteocephalus cabrerai and contained four species, one of which is Osteocephalus buckleyisensu stricto and three are new. Two of the new species are shared between Ecuador and Peru (Osteocephalus vilmaesp. n. and Osteocephalus cannatellaisp. n.) and one is distributed in the Amazon region of southern Peru (Osteocephalus germanisp. n.) We discuss the difficulties of using morphological characters to define species boundaries and propose a hypothesis to explain them.

ALLOZYME AND ADVERTISEMENT CALL VARIATION IN THE TÚNGARA FROG, PHYSALAEMUS PUSTULOSUS.

We analyzed variation in advertisement calls and allozymes in 30 populations along a 5000-km transect throughout most of the range of the túngara frog, Physalaemus pustulosus. All 12 call variables measured show significant differences among populations despite the importance of the advertisement call in species recognition. Some call variables exhibited clinal variation, whereas most others differed between the two major allozyme groups that have invaded Panama at different times, perhaps 4-4.5 million yr apart. Call variables that primarily affect discrimination among conspecifics tended to exhibit greater variation than call variables that are crucial for species recognition. The proximate mechanism of production underlying a call variable, however, is a better predictor of its variation. Contrary to predictions of some sexual selection models, call variation exhibits predictable patterns of geographical variation, although a substantial portion of variation among populations is not explained by geographic position. Although allozymes, calls, and geography usually covary, closer populations can have more similar calls independent of allozyme similarity.

REINFORCEMENT AND REPRODUCTIVE CHARACTER DISPLACEMENT IN GASTROPHRYNE CAROLINENSIS AND G. OLIVACEA (ANURA: MICROHYLIDAE): A REEXAMINATION.

Tape-recorded advertisement calls of Gastrophryne carolinensis and G. olivacea, obtained in Texas and southern Louisiana, were analyzed by means of an analogue audiospectrograph. Samples were grouped into four areas: allopatric and sympatric for G. carolinensis, and combined adjacent allopatric/shallow sympatric, and sympatric for G. olivacea. Three attributes of the advertisement call (call duration, pulse rate, and dominant frequency) were investigated, with water temperature at the calling site as the independent variable. Values for dominant frequency do not overlap between species, across the full range of recording temperatures, and those of sympatric G. carolinensis are displaced away from those of both groups of G. olivacea (which are very similar)-thus indicating a pattern of geographic variation consistent with reproductive character displacement. There is considerable overlap in the values for duration and for pulse rate of each species when considered alone, but there is only slight overlap of the scatters of points for the pairs of values. For both species, no consistent patterns of correlation were detected between the three attributes of the call and the snout-vent length of the emitter, thus reducing the likelihood that the divergence in calls is due to pleiotropic effects of body size.