The otic region of Doleserpeton (Temnospondyli) and its implications for the evolutionary origin of frogs

There is increasing evidence that the Palaeozoic temnospondyl amphibians had a frog‐like tympanic hearing system. For this reason, the otic region of Doleserpeton is described and compared with modern anurans. The otic capsules are expanded laterally and ventrally relative to other temnospondyls. The opisthotic has a bulbous ventral region resembling the ventrolateral ledge in modern frogs. Two lateral processes are located on the paroccipital process. Comparison with the condition in modern anurans with a tympanic hearing system shows that this may have been the attachment site for the tympanic annulus. Parts of the osseous labyrinth are also described. The inner ear shows numerous features resembling the condition found in frogs. These include strong evidence for the presence of a lissamphibian‐type perilymphatic duct most closely resembling that of anurans. This is the first time such a perilymphatic system has been described in any Palaezoic form. The posterior part of the braincase shows a jugular foramen closely associated with the perilymphatic foramen, as in anurans. Although the distribution of these traits among other temnospondyl groups remains little known, the sum of the evidence points to affinities between anurans and temnospondyls, and adds to the evidence for a close relationship between anurans and the Permian amphibamid Doleserpeton. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 738–751.     

Phylogenetic interrelationships of the barbets (Aves: Capitonidae) and toucans (Aves: Ramphastidae) based on morphology with comparisons to DNA-DNA hybridization

A phylogenetic analysis of the interrelationships of the barbets (Capitonidae) and the toucans (Aves: Ramphastidae, Superfamily Ramphastoidea) is presented. Thirty-two morphological characters from the literature and independent osteological observations were analysed. Character polarity was determined by outgroup comparison to the Picidae, Indicatoridae, Galbulidae, Bucconidae and Coraciiformes. Four alternative phylogenetic hypotheses were compared: (1) the overall most parsimonious morphological phylogeny, (2) the most parsimonious morphological phylogeny in which the capitonids and ramphastids were hypothesized as monophyletic sister groups, and (3) and (4) the most parsimonious hypotheses for the evolution of the morphological characters within two proposed DNA-DNA hybridization phylogenies of the ramphastoids. The analysis focused on the higher level relationships of ramphastids and capitonids and interrelationships among capitonid genera. Two cladistic analyses were performed using 26 phylogenetically informative characters, and the PAUP and CONTREE computer alogorithms. The most parsimonious morphological phylogeny required fewer character changes and had a lower consistency index than any of the alternative hypotheses but congruence between the most parsimonious phylogeny and the second, revised DNA-DNA hybridization hypothesis was very high. Based on these results the monophyly of the Capitonidae is rejected. The ramphastids and the Neotropical capitonids form a well corroborated clade within the pantropical ramphastoid radiation. Neither the African, Asian nor New World capitonids is monophyletic. The genus Trachyphonus is the sister group to all other capitonids and ramphastids. The sister group to the ramphastids is the genus Semnornis. The interrelationships of the Old World capitonids excluding Trachyphonus are not completely resolved by these morphological data but one of the alternative phylogenetic resolutions is presented as a preliminary hypothesis. The clades in this resolved phylogeny are diagnosed and the palaeontology and biogeography of the ramphastoids arc-reviewed in light of this new evidence. A phylogenetic classification is proposed in which the Capitonidae is rejected and the capitonids and ramphastids are placed in seven subfamilies of the Ramphastidae.     

The amphibian octavo-lateralis system and its regressive and progressive evolution

The phylogenetic and ontogenetic changes in the octavolateralis system of sarcopterygian fish and tetrapods, presumed to be important for the formation of an amphibian auditory system, are reviewed. The lateral line system shows rudimentation of lines and loss of ampullary electroreceptors in many amphibians; in some amphibians it never develops. The metamorphic changes of the lateral-line system show different patterns in the different amphibian lineages with metamorphic retention in most urodeles and metamorphic loss in most anurans. The multitude of both ontogenetic and phylogenetic changes of the lateral line system among amphibians do exclude any prediction as to how this system might have changed in ancestral amniotes. The most important auditory epithelium of the tetrapod inner ear, the basilar papilla, seems to be primitively present in all tetrapods and Latimeria. In two amphibian lineages there is a trend towards rudimentation and loss of the basilar papilla. Only in the third order, the anurans, a tympanic ear develops and the inner ear shows a progressive evolution of the auditory epithelia. Together with the known differences in the periotic labyrinth of amphibians and amniotes, this scenario suggests a parallel evolution of the amniotic and anuran auditory periphery. All mechanoreceptive hair cells of the lateral line system and the inner ear appear to receive a common and bilateral efferent innervation. Among amphibians this pattern is represented only in some urodeles, whereas anurans show a derived pattern with loss of a bilateral component and presumably also of a common neuromast/inner ear component. Changes in the rhombencephalic nuclei which receive octavo-lateralis afferent fibers show a trend towards development of auditory nuclei only in the anuran lineage. The phylogenetic appearance of an auditory nucleus in this lineage coincides with the complete absence of formation of ampullary electroreceptors. In contrast, the earlier claim of a correlation between a metamorphic loss of the lateral line system and the formation of an auditory nucleus is not supported by more recent data: an auditory nucleus develops in anurans already prior to metamorphosis and is present in all anurans even when they retain the neuromast system. In anurans with a metamorphic loss of the neuromasts, the second order neurons degenerate as well. This independence of the auditory and the second order lateral line nuclei is further substantiated by their separate projection to other brain areas, like the torus semicircularis of the midbrain, and their functional properties.(ABSTRACT TRUNCATED AT 400 WORDS)     

Large odorous frogs (<Emphasis Type="Italic">Odorrana graminea</Emphasis>) produce ultrasonic calls

We present the first data on the vocalizations of large odorous frogs (Odorrana graminea, previously Odorrana livida), from southern China. The males produce diverse broadband signals most of which contain ultrasonic harmonics. Six basic call-types were identified based on the number of call notes, fundamental frequency, call/note duration, frequency modulation patterns and spectral composition. O. graminea is one of only a few non-mammalian vertebrates able to detect ultrasound, but its tympanic membranes are not recessed. These results should stimulate further studies to provide new insights into the mechanisms underlying high-frequency communication in anurans.     

Development of hearing in vertebrates with special reference to anuran acoustic communication

Amphibians, specially anurans, are excellent model systems for studying acoustic communication. After hatching, anurans exist in two forms; these have two distinct mode of sound perception. Aquatic larvae are perceptive to waterborne sound stimuli; then, following metamorphosis, as terrestrial adults, perceptive to airborne sound stimuli. Added to this, the metamorphosing tadpole presents an equally interesting study as it could recapitulate the events which occurred during the evolution of hearing in vertebrates at the lime of the transition from aquatic to terrestrial life. Metamorphosis entails the loss of a prominent aquatic sensory system—the lateral line system—and the simultaneous gain of another, the inner ear, along with the coevolution of the tympanic middle ear, a basilar papilla and a periotic labyrinth in the inner ear. Another interesting feature is that anurans are believed to be the first terrestrial vertebrates to use vocalization as a part of their reproductive behaviour. Vocal communication plays an important role in behaviour, ranging from territorial defense to reproduction, and calls are classified according to the particular behaviors that they subserve. Adult male anurans produce a species-specific mating call which is used to attract conspecific females dung their mating season, and this call serves as a mechanism in maintaining reproductive isolation from other sympatric species.     

Systematics of the genus Cidaria Treitschke (Lepidoptera, Geometridae, Larentiinae)

The genus Cidaria Treitschke is revised. Eight species of the genus which occur widely in the Palaearctic and northern India, are recognized, of which one, Cidaria luteata sp. nov. , is described as new while two subspecies of C. fulvata (Forster) are elevated to species, C. nugata Felder stat. rev., and C. distinctata Staudinger stat. nov. C. ochripennis Prout is proposed as a junior synonym of C. ochreata Staudinger. C. deletaria Hampson is excluded from the genus. All species of Cidaria and their genitalia are described and illustrated. The cladistic analysis of these eight species of Cidaria is carried out using two data matrices, the first comprising morphological characters alone and the second morphological and distributional characters. The most parsimonious cladogram of Cidaria is selected, and its monophyly defined. The character analysis shows that some distribution characters contribute to resolving the in group node. The choice of multiple trees is discussed.     

The blood parasites of anurans from Costa Rica with reflections on the taxonomy of their trypanosomes

During May 1997, specimens of 7 species of anurans, that included 5 Phrynohyas venulosa Laurenti, 5 Rana forreri Boulenger, 7 Rana vaillanti Brucchi, 6 Eleutherodactylus fitzingeri Schimdt, 4 Smilisca baudinii Duméril and Bibron, 1 Leptodactylus melanonotus, and 3 Bufo marinus Linneaus, from the Guanacaste Conservation Area, Costa Rica were examined for blood parasites. Their hematozoan fauna included intraerythrocytic and intraleukocytic icosahedral viruses, a rickettsia (Aegyptianella sp.), 2 species of Hepatozoon, Lankesterella minima, 2 unknown species of apicomplexans, 9 morphologically distinct types of trypanosomes, and 2 species of microfilariae. Rana vaillanti, the most aquatic species of frog, harbored the most species of parasites. Recent evidence indicates that morphological changes in the highly pleomorphic trypanosomes of anurans from different geographical regions have not kept pace with biochemical (isozyme) and molecular (DNA sequence) changes. Describing new species based solely on bloodstream trypomastigotes is discouraged. Additional criteria described herein should be applied when naming new species of anuran trypanosomes.     

Revisiting the contribution of larval characters to an analysis of phylogenetic relationships of basal anurans

The higher-level relationships of anurans have been explored by numerous studies, producing a variety of hypotheses. The relationships of the basal anurans ('archaeobatrachians') are, however, poorly known . In part, this may be because the adult morphology of basal anurans is derived and therefore may not provide suitable phylogenetic signal. Recently, several authors have shown the phylogenetic utility of information derived from anuran larvae. In this paper we conduct separate and combined analyses of anuran relationships based on adult and larval morphology. Our combined results suggest that anurans form two major clades − the pipoids and all other frogs. Evidence also suggests that, taken together, Neobatrachia and Pelobatoidea form a monophyletic group. We discuss support for various groupings as shown by the different data sets. We also comment on the consequences of our phylogenetic hypothesis for the interpretation of reduction of vertebral elements and evolution of Orton's tadpole types in anurans.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society, 2003, 139 , 129−155.     

Comparative ultrastructural study of normal and grafted skin in the frog,Rana pipiens,with special reference to neuroepithelial connections

Summary Recent investigations have suggested specific differences in back and belly skin in anurans which appear to influence the quality of reflex responses obtained from various areas of the animals body. The present investigation represents a comparative morphological study of back and belly skin in control and skin-graftedRana pipiens, with special regard to the neuroepithelial relationships. A distinct difference in pigmentation of back and belly skin was observed. Intra-epithelial Merkel cells were present in all skin samples studied. The origins of the numerous unclassifiable cells in the Merkel cell region are discussed in relation to a presumed coordinating function of the Merkel cell during epithelial differentiation. Epitheliomesenchymal interactions were observed in the richly innvervated dermal regions. Two types of morphologically different intra-epithelial nerve endings were observed. These observations are discussed in relation to earlier observations on vertebrate skin and in relation to misdirected reflex responses obtained in skin-grafted anurans.     

Molecular phylogeny of the genus Dahlia Cav. (Asteraceae, Heliantheae-Coreopsidinae) using sequences derived from the internal transcribed spacers of nuclear ribosomal DNA

Sequences of the internal transcribed spacers (ITS) of the 18S-26S nuclear ribosomal DNA were used to resolve phylogenetic relationships among 13 species of the genus Dahlia . Phylogenetic reconstruction from both parsimony analysis and pairwise distance data produced a single tree in which four clades could be distinguished, these broadly separating the species into the different chromosome numbers found in Dahlia ( x = 16, x = 17 and x = 18). Although species based on x = 16 were divided into two clades, there is the suggestion that this separation may be artificial and that these should be combined as a single clade. The phylogeny produced here has been discussed with reference to the existing taxonomic treatment of the genus. It currently appears that the genus Dohlia is monophyletic and it is evident that the different basic chromosome numbers withim the dysploid series in Dahlia have evolved only once. This proposed monophyly of the genus coupled with the successful production of hybrids withim the dysploid series (2 n = 32, 2 n = 34 and 2 n = 36) suggests that the species of Dahlia have evolved from a common ancestor.     

Comparative osteology and relationships of the Umbridae (Pisces: Salmoniformes)

The phylogenetic relationships of the five living species of umbrids are examined through a comparative osteological study based on a series of cleared-and-stained specimens of each species. The Umbridae appear to be strictly monophyletic. In addition, for 45 characters, the outgroup Esox shares one character-state with at least one umbrid species, while two or more umbrid species share the other state. Assuming that the states shared with Esox are primitive for the Umbridae, the hypothesis that Dallia is more closely related to Umbra than to Novumbra is supported by compatibility and Wagner-tree analysis of the data. Thirteen derived characters are shared by Dallia and Umbra , 20 more are shared by the three species of Umbra , and four more by U. limi and U. pygmaea. Eight characters are considered likely to have undergone reversal or parallel evolution.     

Evolution of the tetrapod ear: an analysis and reinterpretation

The dominant view of tetrapod otic evolution–the “standard view”–holds that the tympanum developed very early in tetrapod history and is homologous in all tetrapods and that the opercular process of the rhipidistian hyomandibula is homologous to the tympanic process of the stapes in lower tetrapods. Under that view, the labyrinthodont amphibians of the Paleozoic are usually considered ancestral to reptiles, and thus the “otic notch” of labyrinthodonts and the tympanum it presumably contained form the starting-point for middle ear evolution in reptiles. Four problems have classically been identified with the standard view: the differing relationships of the internal mandibular branch of N. VII (chorda tympani) to the processes of the stapes in amniotes and anurans; the differing orientations of the stapes in key fossil and living groups; the location of the tympanum in early fossil reptiles; and the transferral of the tympanum, during the origin of mammals, from the stapes to the articular bone of the lower jaw. An examination of these problems and of the solutions proposed under the standard view reveals the ad hoc, and therefore unsatisfactory, nature of the proposed solutions. To organize and review alternative hypotheses of otic evolution an analytical table is constructed, using three characters (tympanic process, Nerve VII, tympanum), each with two possible states. A total of eight hypotheses about middle ear evolution are possible under this system, one of which is the standard view. The seven “non-standard” hypotheses, only five of which have been argued in the literature, are briefly examined. Six of the “non-standard” hypotheses appear unattractive for various reasons, including reliance on ad hoc arguments. The seventh was first proposed by Gaupp in 1898. It is today almost universally ignored but apparently largely for historical rather than scientific reasons. This hypothesis, her called the “alternative view”, appears to rest on assumptions equally as plausible as those of the standard view. Moreover, it offers a solution of the problems associated with the standard view without, apparently, raising any similarly serious problems. This paper compares the standard and alternative views of middle ear evolution in detail. Comparison proceeds on two levels. On one level, they are compared in terms of the hypotheses of phyletic tetrapod relationships each promotes and how strongly each supports its hypothesis. Both views promote the same hypothesis of tetrapod relationships. The alternative view is the more parsimonious, but the difference is not considered sufficient to provide a choice. On another level, the two views are compared in terms of their implications for: (1) the evolution of relative and absolute auditory perceptive ability; (2) the origin of reptiles; (3) the evolution of the suspensorium and cranial kinesis; and (4) the origin and evolution of recent amphibians. The nature of the data required for a test of the implications of the two views is specified in each case. Where data are available. the alternative view is consistent and the standard view is inconsistent with these data. We conclude that the alternative view is the preferable hypothesis of middle-ear evolution. This conclusion implies the following: the tympanic membranes and the tympanic processes of the stapes in recent mammals, reptiles + birds. and frogs. are not homologous; the evolution of “special periotic systems” in the ancestors of amphibians and amniotes were independent events and preceded the evolution of tympanic membranes; the amphibian tympanic membrane. probably including that of labyrinthodonts. is not ancestral to that of amniotes. and that labyiinthodonts with an otic notch are not suitable as amniote ancestors; the stapes of early reptiles functioned primarily as part of the jaw suspension rather than in hearing; the mechanisms and abilities of sound perception in recent tetrapods are likely to be diverse rather than forming parts of a cline; and the lack of a tympanum in Gymnophiona and Caudata may be a retention of a primitive condition.     

Tympanic sound radiation in the bullfrog Rana catesbeiana

Members of the Rana catesbeiana clade display sexually dimorphic eardrums. In this species assemblage the eardrum of males can be 50% larger than in females of the same body size. There has been, however, no apparent functional explanation for this dimorphism. Measurements of the acoustical coupling (transfer function) of internally generated sound to the enlarged eardrum of male bullfrogs (R. catesbeiana) show distinct energy peaks coincident with those observed in the spectral envelopes of the release and mating calls. Moreover, when the tympanic membranes are artificially damped the spectrum of the release call is drastically altered and the total amount of power radiated decreases substantially. These observations point to a previously unsuspected role for the ears in the sound broadcasting process of the bullfrog and possibly other anurans with similarly modified tympanic membranes. Accepted: 19 July 1997     

The biology of the Pe-ret' Toad, Otophryne robusta- (Microhylidae), with special consideration of its fossorial larva and systematic relationships

Adult and larval Otophryne robusta- were collected in Colombia, South America. Although this genus has been assigned to the family Microhylidae, a variety of both adult and larval features distinguish it from all other microhylids. The adult is a diurnally active leaf mimic with complex agonistic behaviours and vocalizations. It tends to walk rather than hop, and does not burrow in captivity. The larva is unique in having minute, dagger-like, keratinized teeth and a sinistral spiracle at the tip of a long tube. The mature larva lives shallowly buried in sand at the bottom of clear, shallow streams. The anatomy of the tadpole was examined for clues to how it burrows into and survives in sand, as well as to the correct phylogenetic association of the genus. Internally, the tadpole is most similar to microhylid tadpoles and has a wealth of oral features (e.g. unperforated internal nares, branchial food traps arranged in crescentic organs, ventral velum divided on the midline, glottis located rostral to the free edge of the ventral velum, large gill filters etc.) that characterize that family. Most, if not all, of the unique features of the Otophryne larva can be associated with its fossorial existence. The arrangement of the muscles suggest that it can actively raise and lower its snout during burrowing and possibly dorsiflex its head on its vertebral column. Although the larval teeth first suggest carnivory, many features indicate that maerophagy is not possible. Stomach contents include a variety of bacteria and micro-organisms, but no fragments of macroscopic animals. We conclude that the Otophryne tadpole is a microphagous suspension feeder and its teeth are more important for keeping sand grains out of its mouth than for grasping prey. Since the O. robusta larva does not burrow deeply, the tip of its spiracular tube probably extends upward above the sandy bottom and into the current. In this position pressure would be lower at the spiracle than at the mouth and a current could be passively drawn (by the Bernoulli effect) through the oral cavity for feeding and respiration. If this is correct then the Otophryne larva is the first fossorial vertebrate known to passively filter-feed. All features of the tadpole, except the keratinized teeth and sinistral spiracle, indicate an affinity of Otophryne-/i with South American microhylids. The tadpole and adult are, however, so unusual as to warrant subfamilial status+ADs- we propose the subfamily Otophryninac for this monotypic genus.     

Comparative features of the surface morphology of the basilar papilla in five families of salamanders (Amphibia; Caudata)

The surface morphology of the basilar recess and papilla was examined in 14 species of newts and salamanders selected from the five families of urodeles (Ambystomatidae, Salamandridae, Hynobiidae, Cryptobranchidae, and Amphiumidae) known to have this end-organ. In this sampling, the general organization of basilar structures is essentially similar across species investigated. The recess forms a tubular diverticulum of the proximal part of the lagena. One wall of the recess is associated with a diverticulum of the intracapsular periotic sac, and an adjacent wall is occupied by the basilar papilla. The papilla contained from as few as five hair cells in specimens of Taricha torosa to over 200 hair cells in Cryptobranchus allegheniensis. In most species, the papilla showed a morphological continuum between tall centrally or distally placed ciliary bundles and short ciliary bundles near the papillar margins. In certain species examined, tall bundles had kinocilia with swellings near their tips. Most forms showed a tendency to have groups of ciliary bundles morphologically polarized either toward or away from the saccule. In Cryptobranchus and Dicamptodon, many bundles had a random orientation. The gross and fine structural features of the basilar complex are compared in urodeles and anurans, and "generalized" features for the amphibian basilar complex are suggested. The basilar complex of Cryptobranchus is interpreted as being most generalized, representing a structural form from which most features of the basilar complex in other urodeles and anurans can be derived.     

Evidence of stochasticity driving anuran metacommunity structure in the Pantanal wetlands

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Wood and stem anatomy of Lardizabalaceae, with comments on the vining habit, ecology and systematics

CARLQUIST, S., 1984. Wood and stem anatomy of Lardizabalaceae, with comments on the vining habit, ecology and systematics. Qualitative and quantitative data, based mostly upon liquid-preserved specimens, are presented for Akebia, Roquila, Decaisnea, Holbodia, Lardizabala, Sinofranchetia and Stauntonia . Because Decaisnea is a shrub whereas the other genera are vines, anatomical differences attributable to the scandent habit can be considered. These include exceptionally wide vessels, a high proportion of vessels to tracheids (or other imperforate trdcheary elements) as seen in transection, simple perforation plates, multiseriate rays which are wide and tall, and pith which is partly or wholly sclerenchymatous. With respect to ecology, two features are discussed: spirals in narrower vessels may relate to adaptation to freezing in the species of colder areas, and crystalliferous sclereids seem adapted in morphology and position to deterrence of phytophagous insects or herbivores. The wood may provide mechanisms for maintaining conduction even if wider vessels are deactivated temporarily by formation of air embolisms. Wood and stem anatomy of Lardizabalaceae compare closely to those of Berberidaceae and of Clematis (Ranunculaceae), as well as to other families of Berberidales. Decaisnea is more primitive than these in having consistently sralariform perforation plates and in having scalariform pitting on lateral walls of vessels. A tentative listing of anatomical features which may correspond to generic limits is given.     

Anuran and pig egg zona pellucida glycoproteins in fertilization and early development

The envelope surrounding the eggs of all animals has many biological functions in fertilization and development. This review focuses on the anuran egg envelope in terms of its biochemistry, biophysics, structural biology and function in sperm-egg interactions and early development (embryo hatching). Egg envelopes from Xenopus laevis are among the most studied of the anurans, and are the central theme of this review. Comparisons of Xenopus laevis envelopes with those of other anurans and with pig egg envelopes are also included. This article presents historical as well as contemporary comparative perspectives on molecular and cellular mechanisms of sperm-egg envelope binding, block to polyspermy, envelope hardening, and hatching.     

Male secondary sexual characteristics, sexual selection, and molecular divergence in fanged ranid frogs of Southeast Asia

A group of Southeast Asian frogs of the family Ranidae are characterized by a suite of unusual secondary sexual characteristics. Males are larger than females, possess bony odontoid processes (fangs) and have enlarged heads. While all members of this clade are sexually dimorphic in this suite of characters, the degree of development of the secondary sexual characteristics in males varies widely across species. One of the most common species, Rana blythii Boulenger, has an extremely broad geographical range. This frog lacks an advertisement call in Borneo but calling has been reported in populations from peninsular Malaysia and Vietnam. This study examines the relationship between the expression of male secondary sexual characteristics, molecular divergence and number of speciation events in widely separated populations of R. blythii , and between R. blythii and other species of closely related fanged frogs belonging to the grunniens species group. Results from morphometric and sequence data indicate that (1) there is no correlation between morphological and molecular distance in these frogs and (2) there is no correlation between morphological distance and the number of speciation events separating taxa. Systematic analysis demonstrates that R. blythii , as currently recognized, is not a single species nor is it monophyletic.     

Anuran occupancy of created wetlands in the Central Appalachians

Evaluating the adequacy of created wetlands to replace the functions of lost natural wetlands is important because wetland mitigation is a major tool used to offset wetland losses. However, measurements such as vegetative cover and presence of wildlife may not provide sufficient evidence that created wetlands are functioning properly and thus examining the ecology of wetland biota such as that of amphibians may be a more useful surrogate for function. The objectives of this study were to compare the occupancy and detection of calling anurans in created wetlands relative to beaver-created wetlands. Five-, 10-min, and broadcast call surveys were performed at 24 wetlands throughout the Central Appalachians once every month from March through August of 2009 and 2010. Occupancy modeling was used to estimate the occupancy and detection of individual species, incorporating relevant environmental variables. The occupancy of anurans did not differ between human-created and beaver (Castor canadensis)-created wetlands. Detection of anurans was largely unaffected by call survey type, but several environmental covariates had a significant effect on the detectability of calling anurans. Our results suggest that the function of providing adequate chorusing habitat for adult anurans is being fulfilled by the created wetlands that we examined.