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Charles Darwin proposed a separate theory of sexual selection, as distinct from his theory of natural selection, to account for adaptations that confer success in finding a mate, which may sometimes be quite the opposite of what is best for survival. Darwin's proposal that females have a sense of beauty and choose mating partners that appear beautiful to them was met with much scepticism. But today we have a rather detailed understanding of what animals find beautiful and why. In this article, I will describe a few very simple experiments performed by Michael J. Ryan, in collaboration with A. Stanley Rand, herpetologist extraordinaire and Merlin D Tuttle of the Bat Conservation International fame, that laid the foundation for our current understanding of the meaning and evolution of beauty. Studying the túngara frog on Barro Colorado Island, a research station of the Smithsonian Tropical Research Institute in Panama, they showed that (1) male túngara frogs can produce both simple calls, consisting of just a whine, or complex calls in which one or more chucks are added to the whine, (2) female túngara frogs have a decided preference to mate with males giving complex calls, (3) males are nevertheless reluctant to add chucks to their calls and generally do so only when they hear other males calling, and (4) the local predatory fringe-lipped bat also has a decided preference to eat males giving complex calls. Male tungara frogs thus face a trade-off between sex and survival. These experiments not only answered the question of why males don't do their best when it comes to singing, but they also set the stage for many more sophisticated investigations that have led to an understanding of how and why natural selection has favoured this particular sexual aesthetic in the frogs and this particular culinary aesthetic in the bats. [ABSTRACT FROM AUTHOR]

Relatively few studies have used experimental manipulations to investigate the mechanics of vocal production in frogs and toads, even though many frogs produce complex signals with multiple components and/or nonlinearities. Modelling approaches can add to empirical studies by illuminating how various components of the vocal system interact to produce communication signals. In this study, we use bond graphs, a lumped-element modelling technique, to explore how the combination of active modulation of vocal production and the passive dynamics of a unique laryngeal structure result in the complex calls produced by males in an anuran model system. The túngara frog (Physalaemus (= Engystomops) pustulosus) produces advertisement calls with a 'whine' and a facultative 'chuck'. Whines are amplitude and frequency modulated. The chuck is characterized by its spectral complexity and appears to contain a period doubling bifurcation resulting in subharmonics. In our model, we focus on how a fibrous mass attached to the vocal cords results in subharmonics in the chuck. Our models suggest that active (neural) modulation of the fibrous mass is not necessary for the transition between the spectral characteristics of the whine and chuck. Rather, it is possible that the vibratory mode of the fibrous mass and thus the vocal cords changes passively as a result of changes in airflow through the system. [ABSTRACT FROM AUTHOR]

Sexual selection driven by mate choice has generated some of the most astounding diversity in nature, suggesting that population-level preferences should be strong and consistent over many generations. On the other hand, mating preferences are among the least repeatable components of an individual animal's phenotype, suggesting that consistency should be low across an animal's lifetime. Despite decades of intensive study of sexual selection, there is almost no information about the strength and consistency of preferences across many years. In this study, we present the results of more than 5,000 mate choice tests with a species of wild frog conducted over 19 consecutive years. Results show that preferences are positive and strong and vary little across years. This consistency occurs despite the fact that there are substantial differences among females in their strength of preference. We also suggest that mate preferences in populations that are primarily the result of sensory exploitation might be more stable over time than preferences that are primarily involved in assessing male quality. [ABSTRACT FROM AUTHOR]

Most models of animal foraging assume that individuals make decisions rationally. One expectation of rationality theory is that preferences should be hierarchical or transitive. For many animals, including humans, making decisions about what to eat can become difficult when various parameters of choices are not positively related to one another and no single option may be considered the best. These complex decisions can result in irrational behaviour, suggesting that the value of any particular option may vary depending on the other available options. Here, we tested whether frog-eating bats, Trachops cirrhosus , are transitive in their preferences for frog calls. These bats attend to several attributes of the calls, such as call amplitude and complexity, when choosing a frog to target in their attack. While acoustic preferences are largely understood in isolation, we have a limited understanding of how bats choose between calls that vary along several important acoustic dimensions. Here, we conducted a series of binary choice experiments manipulating amplitude and complexity both separately and together to evaluate whether these bats make transitive foraging decisions. We found that bats are transitive in their rankings of calls based on these acoustic dimensions, with preferences reflecting a trade-off between both amplitude and complexity. Overall, our results add support to the notion that bats' preferences obey the general principles of rational behaviour and support the use of optimality approaches to the study of animal behaviour and decision making. • Animal foraging models assume hierarchical preferences, but evidence is mixed. • Frog-eating bats showed linear preferences for calls that varied in amplitude. • Bats preferred calls with chucks and calls with more chucks up to a threshold. • When chucks and amplitude were varied, bats showed transitive preferences. [ABSTRACT FROM AUTHOR]

Vocalizations play a critical role in mate recognition and mate choice in a number of taxa, especially, but not limited to, orthopterans, frogs, and birds. But receivers can only recognize and prefer sounds that they can hear. Thus a fundamental question linking neurobiology and sexual selection asks—what is the threshold for detecting acoustic sexual displays? In this study, we use 3 methods to assess such thresholds in túngara frogs: behavioral responses, auditory brainstem responses, and multiunit electrophysiological recordings from the midbrain. We show that thresholds are lowest for multiunit recordings (ca. 45 dB SPL), and then for behavioral responses (ca. 61 dB SPL), with auditory brainstem responses exhibiting the highest thresholds (ca. 71 dB SPL). We discuss why these estimates differ and why, as with other studies, it is unlikely that they should be the same. Although all of these studies estimate thresholds they are not measuring the same thresholds; behavioral thresholds are based on signal salience whereas the 2 neural assays estimate physiological thresholds. All 3 estimates, however, make it clear that to have an appreciation for detection and salience of acoustic signals we must listen to those signals through the ears of the receivers. [ABSTRACT FROM AUTHOR]

Advertisement signals can convey information about a sender's characteristics, such as body size. The reliability of signals, however, can be reduced when signal production is partially dependent on the environment. Here, we assess the effect of display-site properties on the production, attractiveness and honesty of sexual signals. We recorded male túngara frogs (Physalaemus pustulosus) while manipulating water levels in order to constrain calling. We found that water level affected male call [ABSTRACT FROM AUTHOR]

The traits of many animal species exhibit individual and sexual differences. Individuals repeatedly receiving a stimulus without harm become habituated to it. However, few studies have been conducted on individual and sexual differences in the process of habituation to unfamiliar food stimuli. Therefore, we hypothesized that individual differences or sexual differences would be observed in reaction to an in-lab food-stimuli presentation of potential prey items (after that "food stimuli"). We tested the hypothesis using the Japanese tree frog Hyla japonica, and conducted statistical analyses of these results. A generalized linear model (GLM) showed individual and sexual differences in time to get used to the food stimuli. Females habituated more rapidly to food stimuli than males. The difference between sexes is discussed in view of two ultimate and one proximate reasons. [ABSTRACT FROM AUTHOR]

Animals are expected to optimize energy intake when choosing between different foraging options. A common explanation for deviations from optimal economic decisions is that there is an imperfect relationship between physical reality and an animal's perceptual processes, which can constrain assessment of profitability. One such phenomenon that is apparently ubiquitous across taxa is proportional processing, where a perceived change in a stimulus is proportional to the change in stimulus magnitude. In this study, we investigated whether proportional processing explains how frog-eating bats, Trachops cirrhosus, discriminate between patches of frog choruses that vary in their number of calling frogs. To test this, we created artificial choruses consisting of one to six calling frogs. In the flight cage, we then tested the preference of bats ( N = 17) with every pairwise combination of chorus size. We found that while bats generally preferred larger choruses, preferences for larger choruses were better explained by the relative, not absolute, differences in chorus sizes. This indicates that T. cirrhosus is perceptually limited in its ability to discriminate between choruses of varying size as the choruses increase in size. Foragers are likely to be less choosy when choosing among larger patches. Highlights • We investigated perceptual limitations in patch size assessment in frog-eating bats. • Frog-eating bats preferred larger choruses, with more calling frogs. • Bats attended to relative, not absolute, differences when choosing larger chorus sizes. [ABSTRACT FROM AUTHOR]

The evolution of elaborate signals can emerge from changes in anatomical signaling structures. In the male túngara frog, a simple advertisement call (‘whine’) can be ornamented facultatively with a suffix (‘chuck’) to produce a more attractive complex call, or ‘whine-chuck’. A fibrous mass (FM1) supported by the vocal cords plays a role in chuck production. Here, we examine the effects of FM1 ablation on a large set of spectral and temporal features of both the whine and chuck and we test the hypothesis that FM1 ablation reduces call attractiveness to females. Both call components were impacted by FM1 ablation, but especially the suffix. The proportion of energy in the suffix’s odd compared to even harmonics diminished markedly as did the relative amplitude, effectively eliminating the chuck percept. FM1 ablation also reduced the whine’s frequency and its rate of frequency modulation. Moreover, post-surgery chucks no longer enhanced the attractiveness of the simple call and whines also appeared to diminish in attractiveness. Together, our results demonstrate that FM1 plays a role in the production of both call components in a way that stimulates the female auditory system and that the growth of FM1 had potentially positive sexual selection implications for the pre-existing simple call. [ABSTRACT FROM AUTHOR]

Minimally invasive methods for estimating hormone concentrations in wild vertebrates offer the opportunity to repeatedly measure behavior and hormone concentrations within individuals while minimizing experimenter interference during sample collection. We examined three steroid hormones (corticosterone, CORT; 17-β estradiol, E 2 ; progesterone, PROG) in túngara frogs ( Physalaemus pustulosus ) using non-invasive water-borne methods. Using solid-phase extraction of water samples and liquid extraction of plasma and homogenate samples, coupled with enzyme immunoassays, we complimented the conventional validation approaches (parallelism, recovery determination) with dose-response assays that incorporated pharmacological challenges with adrenocorticotropic hormone (ACTH) and human chorionic gonadotropin (HCG). We also compared steroid concentrations in water to those observed in plasma and whole body homogenates. Lastly, we identified the constituent steroids in each sample type with a panel targeting 30 steroid species using high performance liquid chromatography-mass spectrometry (HPLC-MS). We found that a 60-min water-bath captures physiologically relevant changes in concentrations of CORT, E 2 and PROG. Peak levels of water-borne CORT were found at approximately 2 h after ACTH injection. Water-borne CORT and E 2 concentrations were positively correlated with their plasma and homogenate equivalents, while water-borne PROG was uncorrelated with homogenate PROG concentrations but negatively correlated with homogenate E 2 concentrations. Together, our findings indicate that sampling water-borne hormones presents a non-invasive and biologically informative approach that will be useful for behavioral endocrinologists and conservation physiologists. [ABSTRACT FROM AUTHOR]

Multimodal communication has received increasing attention in recent years. While much is understood about how intended receivers (such as potential mates) respond to multimodal displays, less is known about how eavesdropping predators perceive and interpret these cues. The male túngara frog, Physalaemus pustulosus , is a neotropical anuran that attracts females with an acoustic call and a dynamically inflating/deflating vocal sac. However, the túngara frog's multimodal courtship display also attracts eavesdropping predators, such as fringe-lipped bats, Trachops cirrhosus . We utilized robotic frog models to expose fringe-lipped bats to multimodal túngara frog courtship displays. The models varied in call amplitude and/or the presence of vocal sac cues. In a two-choice test, we show that fringe-lipped bats more often attack higher-amplitude calls. Additionally, coupling the inflating vocal sac cues to the lower-amplitude frog call increased the probability that a bat would attack this less attractive call. Previous studies have demonstrated that vocal sac cues do not increase the attractiveness of low-amplitude calls to female P. pustulosus . Thus, although natural selection, through the bats, and sexual selection, through the female frogs, exert counter-selection forces on the male's sexual display, the strength of these forces are not symmetrical. We discuss possible explanations for why this might be the case. This study underlines the importance of understanding the contribution of both intended and unintended receivers on signal evolution, and it helps explain how selection pressures might vary across sensory modalities. [ABSTRACT FROM AUTHOR]

Multimodal signaling is common in communication systems. Depending on the species, individual signal components may be produced synchronously as a result of physiological constraint (fixed) or each component may be produced independently (fluid) in time. For animals that rely on fixed signals, a basic prediction is that asynchrony between the components should degrade the perception of signal salience, reducing receiver response. Male túngara frogs, Physalaemus pustulosus, produce a fixed multisensory courtship signal by vocalizing with two call components (whines and chucks) and inflating a vocal sac (visual component). Using a robotic frog, we tested female responses to variation in the temporal arrangement between acoustic and visual components. When the visual component lagged a complex call (whineþchuck), females largely rejected this asynchronous multisensory signal in favor of the complex call absent the visual cue. When the chuck component was removed from one call, but the robofrog inflation lagged the complex call, females responded strongly to the asynchronous multimodal signal. When the chuck component was removed from both calls, females reversed preference and responded positively to the asynchronous multisensory signal. When the visual component preceded the call, females responded as often to the multimodal signal as to the call alone. These data show that asynchrony of a normally fixed signal does reduce receiver responsiveness. The magnitude and overall response, however, depend on specific temporal interactions between the acoustic and visual components. The sensitivity of túngara frogs to lagging visual cues, but not leading ones, and the influence of acoustic signal content on the perception of visual asynchrony is similar to those reported in human psychophysics literature. Virtually all acoustically communicating animals must conduct auditory scene analyses and identify the source of signals. Our data suggest that some basic audiovisual neural integration processes may be at work in the vertebrate brain. [ABSTRACT FROM AUTHOR]

The adaptive specialization hypothesis states that sex differences in cognition are shaped by differences in cognitive demands to solve ecological problems. While it is widely accepted that female mate choice can lead to the evolution of exaggerated male traits, mate choice might also select for different cognitive abilities in males and females. In the túngara frog, Physalaemus (= Engystomops ) pustulosus , males call from a fixed position in breeding ponds while females visit multiple males before returning to the preferred mate. Thus, we predicted that females have better place memory than males. We tested this prediction in a place-learning task in which the rewarded arm of a maze was associated with a visual cue. We found that females were able to use the visual cue to solve the task while males were not, even though both males and females could discriminate the cues in an optomotor test. In contrast, males attempted to solve the task using egocentric cues (remember body-turn direction) in spite of the fact that our training procedure interrupted their use of such cues. Finally, we found that males and females had similar motivation to solve the task but females showed a greater ability to inhibit incorrect responses, leading to improved learning. Our finding that females could use a visual cue to remember locations in space is consistent with the idea that place memory could improve sequential mate assessment in túngara frogs. [ABSTRACT FROM AUTHOR]

Studies on the consequences of urbanization often examine the effects of light, noise, and heat pollution independently on isolated species providing a limited understanding of how these combined stressors affect species interactions. Here, we investigate how these factors interact to affect parasitic frog-biting midges ( Corethrella spp.) and their túngara frog ( Engystomops pustulosus) hosts. A survey of túngara frog calling sites revealed that frog abundance was not significantly correlated with urbanization, light, noise, or temperature. In contrast, frog-biting midges were sensitive to light pollution and noise pollution. Increased light intensity significantly reduced midge abundance at low noise levels. At high noise intensity, there were no midges regardless of light level. Two field experiments controlling light and noise levels to examine attraction of the midges to their host and their feeding behavior confirmed the causality of these field patterns. These findings demonstrate that both light and noise pollution disrupt this host-parasite interaction and highlight the importance of considering interactions among species and types of pollutants to accurately assess the impacts of urbanization on ecological communities. [ABSTRACT FROM AUTHOR]

The neuropeptide arginine vasotocin (AVT) promotes sexual advertisement and influences vocalization structure in male anuran amphibians. In the present study, we used wild túngara frogs ( Physalaemus pustulosus) to investigate the effects of AVT on phonotaxis in males and females-thereby controlling for potential task differences between the sexes. Using a combined within- and between-subjects design, we showed that acoustic choice behavior in female frogs is not influenced by injection per se (vehicle) or by AVT. Latency to choice in females, however, tends to decrease after AVT injection, supporting the hypothesis that AVT promotes female sexual arousal. In contrast, male choice behavior and latencies are negatively impacted by injection (vehicle) but rescued to pre-injection levels if administered with AVT. The sexes differed in area restricted searching (ARS) following choice-a measure of locomotor perseverance-with females but not males exhibiting ARS. AVT did not influence ARS behavior but ARS frequency was positively associated with the attractiveness of the acoustic stimulus. Finally, we showed that a female's latency behavior is correlated with her partner's behavior. Collectively we show that AVT promotes phonotaxis in both sexes in a dimorphic manner-a result that is consistent with sex differences in the neural vasotocin system. [ABSTRACT FROM AUTHOR]

Perceptually, grouping sounds based on their sources is critical for communication. This is especially true in túngara frog breeding aggregations, where multiple males produce overlapping calls that consist of an FM 'whine' followed by harmonic bursts called 'chucks'. Phonotactic females use at least two cues to group whines and chucks: whine-chuck spatial separation and sequence. Spatial separation is a primitive cue, whereas sequence is schema-based, as chuck production is morphologically constrained to follow whines, meaning that males cannot produce the components simultaneously. When one cue is available, females perceptually group whines and chucks using relative comparisons: components with the smallest spatial separation or those closest to the natural sequence are more likely grouped. By simultaneously varying the temporal sequence and spatial separation of a single whine and two chucks, this study measured between-cue perceptual weighting during a specific grouping task. Results show that whine-chuck spatial separation is a stronger grouping cue than temporal sequence, as grouping is more likely for stimuli with smaller spatial separation and non-natural sequence than those with larger spatial separation and natural sequence. Compared to the schema-based whine-chuck sequence, we propose that spatial cues have less variance, potentially explaining their preferred use when grouping during directional behavioral responses. [ABSTRACT FROM AUTHOR]

Disentangling the influence of multiple signal components on receivers and elucidating general processes influencing complex signal evolution are difficult tasks. In this study we test mate preferences of female squirrel treefrogs Hyla squirella and female túngara frogs Physalaemus pustulosus for similar combinations of acoustic and visual components of their multimodal courtship signals. In a two-choice playback experiment with squirrel treefrogs, the visual stimulus of a male model significantly increased the attractivness of a relatively unattractive slow call rate. A previous study demonstrated that faster call rates are more attractive to female squirrel treefrogs, and all else being equal, models of male frogs with large body stripes are more attractive. In a similar experiment with female túngara frogs, the visual stimulus of a robotic frog failed to increase the attractiveness of a relatively unattractive call. Females also showed no preference for the distinct stripe on the robot that males commonly bear on their throat. Thus, features of conspicuous signal components such as body stripes are not universally important and signal function is likely to differ even among species with similar ecologies and communication systems. Finally, we discuss the putative information content of anuran signals and suggest that the categorization of redundant versus multiple messages may not be sufficient as a general explanation for the evolution of multimodal signaling. Instead of relying on untested assumptions concerning the information content of signals, we discuss the value of initially collecting comparative empirical data sets related to receiver responses [Current Zoology 57 (2): 153–161, 2011].

The potential for ornament evolution in response to sexual selection rests on the interaction between the permissiveness or selectivity of female preferences and the constraints on male development of signaling related traits. We investigate the former by determining how latent female preferences either exaggerate the magnitude of current traits (i.e. elaborations) or favor novel traits (i.e. innovations). In túngara frogs, females prefer complex mating calls (whine-chucks) to simple calls (whine only). The whine is critical for mate recognition while the chuck further enhances the attractiveness of the call. Here we use a combination of synthetic and natural stimuli to examine latent female preferences. Our results show that a diversity of stimuli, including conspecific and heterospecific calls as well as predator-produced and human-made sounds, increase the attractiveness of a call when added to a whine. These stimuli do not make simple calls more attractive than a whine-chuck, however. In rare cases we found stimuli that added to the whine decrease the attractiveness of the call. Overall, females show strong preferences for both elaborations and innovations of the chuck. We argue that the emancipation of these acoustic adornments from mate recognition allows such female permissiveness, and that male constraints on signal evolution are probably more important in explaining why males evolved their specific adornment. Experimentally probing latent female preferences for stimuli out of the species’ range is a useful means to gain insights about the potential of female choice to influence signal evolution and thus the astounding diversity in male sexually-selected traits [Current Zoology 56 (3): 343–357, 2010].

Perceptual-biases are important for understanding an animal's natural history, identifying potential ecological traps, and for developing effective means to monitor individuals and populations. Despite research demonstrating anurans having a positive phototactic response towards blue colors, we do not yet understand if color cues are used functionally beyond sexual selection. The aim of our study was to determine if color cues are used in selecting microhabitat, and if anuran's blue-positive phototactic response could increase selection of artificial PVC refugia used to monitor cryptic camouflaging anuran species. We captured 32 Cope's Gray Treefrogs and placed them in mesh enclosures with three PVC tubes painted blue, brown, and white. Concurrently, we placed blue, brown, or unpainted white PVC tubes in stratified arrays around a treefrog breeding pond, and counted the number of occasions treefrogs occupied different colored PVC tubes. In the confined choice experiment, treefrogs selected blue tubes (48.3%) significantly more often than brown (28.5%) or white (23.2%) tubes. Our field experiment mirrored these findings (52.0% of capture events in blue, 29.0% in brown, and 19.0% in unpainted white tubes). Our results suggest color influences Cope's Gray Treefrog microhabitat selection, and they utilize color vision when choosing refugia. We demonstrate simple, small changes based on perceptual-biases can induce behaviors that may in turn have large impacts on sampling techniques used in monitoring and inventorying. Incorporating non-traditional physiological measures into animal inventorying and monitoring programs can be used in the future to improve conservation efforts. [ABSTRACT FROM AUTHOR]

The world is a noisy place, and animals have evolved a myriad of strategies to communicate in it. Animal communication signals are, however, often multimodal; their components can be processed by multiple sensory systems, and noise can thus affect signal components across different modalities. We studied the effect of environmental noise on multimodal communication in the túngara frog (Physalaemus pustulosus). Males communicate with rivals using airborne sounds combined with call-induced water ripples. We tested males under control as well as noisy conditions in which we mimicked rain- and wind-induced vibrations on the water surface. Males responded more strongly to a multimodal playback in which sound and ripples were combined, compared to a unimodal sound-only playback, but only in the absence of rain and wind. Under windy conditions, males decreased their response to the multimodal playback, suggesting that wind noise interferes with the detection of rival ripples. Under rainy conditions, males increased their response, irrespective of signal playback, suggesting that different noise sources can have different impacts on communication. Our findings show that noise in an additional sensory channel can affect multimodal signal perception and thereby drive signal evolution, but not always in the expected direction. [ABSTRACT FROM AUTHOR]

Many animal communication systems have evolved signal flexibility depending on environmental conditions. A common strategy of vocal communication is to increase amplitude in response to increasing noise levels. This phenomenon, known as the Lombard effect, is a widespread trait among mammals and birds. Anurans are a major group with many species that rely heavily on acoustic signals for sexual communication. Although these species often communicate in noisy environments, the presence of the Lombard effect in frogs remains unclear. We exposed male túngara frogs (Physalaemus pustulosus) to different noises with and without playback of conspecific calls. Males increased call amplitude, call rate, and call complexity in response to low-frequency noise (overlapping the species' call range) but not to high-frequency (nonoverlapping) noise. Vocal amplitude increased linearly with noise level demonstrating that túngara frogs exhibit the Lombard effect, and we discuss why different frog species may differ in their control over vocal amplitudes. Furthermore, we found the overall effect of noise to be similar to the effect of conspecific call playback. We speculate that vocal amplitude control may have evolved primarily as a response to increased competition at the cocktail party, similar to the way humans raise their voice when in a heated debate, and subsequently as a strategy to deal with background noise more generally. [ABSTRACT FROM AUTHOR]

Estradiol plays an important role in mediating changes in female sexual behavior across reproductive cycles. In the túngara frog [Physalaemus (=Engystomops) pustulosus], the relationship between gonadal activity and female sexual behavior, as expressed by phonotaxis, is mediated primarily by estradiol. Estradiol receptors are expressed in auditory and motivational brain areas and the hormone could serve as an important modulator of neural responses to conspecific calls. To better understand how estradiol modifies neural responses to conspecific social signals, we manipulated estradiol levels and measured expression of the immediate early gene egr-1 in the auditory midbrain, thalamus and limbic forebrain in response to conspecific or heterospecific calls. We found that estradiol and conspecific calls increased egr-1 expression in the auditory midbrain and limbic forebrain, but in the thalamus, only conspecific calls were effective. In the preoptic area, estradiol enhanced the effect of the conspecific call on egr-1 expression, suggesting that the preoptic area could act as a hormonal gatekeeper to phonotaxis. Overall, the results suggest that estradiol has broad influences on the neural circuit involved in female reproduction, particularly those implicated in phonotaxis. [ABSTRACT FROM AUTHOR]

Signal detection, recognition, and localization are hampered when multiple signalers coincide in time and space, a problem known as 'cocktail party effect'. In many taxa, senders utter complex calls consisting of two or more elements which often vary in the ease with which they can be assessed in different signaling environments. Receivers' selective attention to different cues may increase the probability of correctly assigning a signal to its source (localization) in face of conspecific interference. Túngara frogs, Physalaemus pustulosus, produce complex calls consisting of an initial whine, followed by zero up to seven broad-banded, amplitude-modulated chucks. Under ideal conditions (without interference or noise), females prefer whines followed by chucks over whines alone, but the preference is not linear; females do not discriminate between whines with one or two chucks. When whines lack chucks, call overlap elicits random responses in females, with no preference for leading calls. In this study, I explored the combined effect of call timing and call complexity on female preferences in a two-choice paradigm-a simplification of the cocktail party scenario. I tested the hypothesis that the effect of call overlap can be reduced when the calls of one of the two rivals have chucks, specifically more chucks than those of the rival. I gave females a choice between whines alone and with chucks (one or two) presented at three time relations (alternated, abutted, and partially overlapped) and two emission orders (whine with less chucks leading and whine with more chucks leading). I found that the preference for one chuck over no chuck was preserved in all the experimental treatments, but when a w + 2chk preceded a w + chk, either overlapped or abutted, a preference existed for the whine with more chucks. Therefore, an interaction between call order and the number of chucks was obtained. The results only partially supported the hypothesis, and call order emerges as an opportunistic component of signaling in P. pustulosus. [ABSTRACT FROM AUTHOR]

Eavesdropping predators sometimes show preferences for certain prey signal variants, yet the ultimate and proximate reasons for such preferences are often unclear. The fringe-lipped bat, Trachops cirrhosus, eavesdrops on the advertisement calls of male túngara frogs, Physalaemus pustulosus, and shows a marked preference for complex (adorned) calls over simple (non-adorned) calls. We hypothesized that this preference stems from perceptual biases in the sensory and/or cognitive systems of T. cirrhosus. To test this hypothesis, we conducted a series of preference experiments in which we presented bats with various modified simple calls, each altered to possess one of the acoustic properties that distinguish complex calls from simple calls. We reasoned that if perceptual bias accounts for the bat's preference for complex calls, then a novel stimulus with similar acoustic properties to the complex call should be attractive as well (i.e., the preference should be permissive). Except for weak evidence suggesting that the longer duration of complex calls could contribute to their greater attractiveness to T. cirrhosus, we did not find any indication that perceptual biases account for this eavesdropper preference. Instead, we suggest that T. cirrhosus developed their preference for call complexity because eavesdropping on complex calls provides greater fitness benefits than eavesdropping on simple calls, for example, because eavesdropping on complex calls may increase probability of prey capture and/or lead to more profitable food patches. [ABSTRACT FROM AUTHOR]

Anuran survival is strongly affected by exposure to high environmental temperatures. However, their upper thermal tolerances vary between species and within developmental stages. The aims of this research were to measure the median lethal temperature (LT50) of three anuran developmental stages (Gosner stages 10, 20, and 25) at a constant thermal regime, and of developing embryos (stage 10) until they became tadpoles (stage 25) exposed to daily peaks of temperatures between 1000 and 1600. Four Colombian species (Emerald-eyed Treefrog, Hypsiboas crepitans (Wied-Neuwied, 1824); Tungara Frog, Engystomops pustulosus (Cope, 1864); Rivero's Toad, Rhinella humboldti (Gallardo, 1965); Emerald Glassfrog, Espadarana prosoblepon (Boettger, 1892)) were used in these experiments. An ontogenetic increase was observed in the upper thermal tolerance from embryos to tadpoles for all species studied. In addition, developing embryos exposed to peak temperatures showed a LT50 fairly close to the mean of the maximum habitat temperatures, particularly in H. crepitans and E. pustulosus that lay egg masses exposed directly to the sun. Environmental temperatures in the microhabitat of species studied showed values remarkably higher than their experimental LT50. Therefore, we postulate that rapid increases in environmental temperatures, as result of global or local changes, might be a critical factor for anuran survival, mainly during the embryonic stages when they are more sensitive to temperature. [ABSTRACT FROM AUTHOR]

Many sexual displays contain multiple components that are received through a variety of sensory modalities. Primary and secondary signal components can interact to induce novel receiver responses and become targets of sexual selection as complex signals. However, predators can also use these complex signals for prey assessment, which may limit the evolution of elaborate sexual signals. We tested whether a multimodal sexual display of the male túngara frog (Physalaemus pustulosus) increases predation risk from the fringelipped bat (Trachops cirrhosus) when compared with a unimodal display. We gave bats a choice to attack one of two frog models: a model with a vocal sac moving in synchrony with a mating call (multisensory cue), or a control model with the call but no vocal sac movement (unimodal cue). Bats preferred to attack the model associated with the multimodal display. Furthermore, we determined that bats perceive the vocal sac using echolocation rather than visual cues. Our data illustrate the costs associated with multimodal signaling and that sexual and natural selection pressures on the same trait are not always mediated through the same sensory modalities. These data are important when considering the role of environmental fluctuations on signal evolution as different sensory modalities will be differentially affected. [ABSTRACT FROM AUTHOR]

In the majority of frogs, males but not females produce vocalizations to attract mates. Sexual selection can influence the evolution of these vocalizations by modifying the frog's morphology. The larynx is the main organ responsible for sound production; thus, it constitutes a target of selection. The goal of this study was to determine qualitatively and quantitatively the sexual differences in the developmental sequence of larynges of Physalaemus pustulosus ( = Engystomops pustulosus). We examined larynges of individuals ranging from recent metamorphs to adults. We provide quantitative measurements of laryngeal area and volume, vocal cords, and fibrous mass 1 (FM1), as well as qualitative descriptions of the developmental origin of vocal cords and FM1. Larynges of small male and female juveniles (less than 16 mm SVL) are similar, both externally and internally. The area of these larynges is no more than 1 mm2; no vocal cords or FM1 are present in either sex at this developmental stage. In individuals larger than 16 mm SVL, there is a marked divergence in relative size of the larynx between the sexes. The male larynx begins to exhibit a strong positive allometry until it reaches a plateau at 24 mm SVL. Also, at 16 mm SVL, the vocal cords and FM1 appear as distinctive outgrowths on the wall of the arytenoid cartilage in male larynges. Female growth rate of the larynx, however, is constant and exhibits negative allometry throughout ontogeny. Female vocal cords and FM1 are barely distinguishable as adults, and never attain the size and most likely neither the mobility as in the adult male larynx. [ABSTRACT FROM AUTHOR]

Animal displays are often perceived by intended and unintended receivers in more than one sensory system. In addition, cues that are an incidental consequence of signal production can also be perceived by different receivers, even when the receivers use different sensory systems to perceive them. Here we show that the vocal responses of male túngara frogs (Physalaemus pustulosus) increase twofold when call-induced water ripples are added to the acoustic component of a rival's call. [ABSTRACT FROM AUTHOR]

The mating calls of male túngara frogs, Physalaemus pustulosus, attract intended (conspecific females) and unintended (eavesdropping predators and parasites) receivers. The calls are complex, having two components: a frequency-modulated 'whine' followed by 0-7 harmonic bursts or 'chucks'. The whine is necessary and sufficient to elicit phonotaxis from females and the chuck enhances call attractiveness when it follows a whine. Although chucks are never made alone, females perceptually bind the whine and chuck when they are spatially separated. We tested whether an unintended receiver with independent evolution of phonotaxis, the frog-eating bat, Trachops cirrhosus, has converged with frogs in its auditory grouping of the call components. In contrast to frogs, bats approached chucks broadcast alone; when the chuck was spatially separated from the whine the bats preferentially approached the whine, and bats were sensitive to whine-chuck temporal sequence. This contrast suggests that although disparate taxa may be selected to respond to the same signals, different evolutionary histories, selective regimes, and neural and cognitive architectures may result in different weighting and grouping of signal components between generalist predators and conspecific mates. [ABSTRACT FROM AUTHOR]

The acquisition of signals used in intrasexual communication is not well studied. To better understand the possible contribution of early experience to the acquisition of intrasexual responses, we reared Neotropical Tungara frogs, Physalaemus pustulosus, in four acoustic environments: 1) conspecific chorus, 2) heterospecific chorus, 3) acoustic isolation, and 4) noise. For the heterospecific chorus, we chose the calls of a congener, P. enesefae, with which P. pustulosus is sympatric in parts of its distribution. We measured the evoked vocal responses (EVR) of these frogs in response to calls of the conspecifics, to calls of the congener, and to calls representing an intermediate between the two species. Male P. pustulosus reared hearing the calls of P. enesefae produced more calls overall and more complex calls in response to the call of P. enesefae. This is consistent with the hypothesis that the EVR of male anurans can be influenced by early experience and is the product of an interaction between genes and the environment. [ABSTRACT FROM AUTHOR]

Female mate choice preferences may be the result of innate factors, experience, or an interaction between the two mechanisms. Understanding the relative contribution and possible interaction between these mechanisms is important for identifying sources of variation in behaviors under sexual selection. In this study, we evaluate the contribution of early experience on the development of mate choice preferences in the Neotropical Tüngara Frog, Physalaemus pustulosus. We reared frogs from tadpole stage through sexual maturity in four acoustic treatments: the first group heard a conspecific chorus of Tüngara Frogs, the second group heard a heterospecific chorus of the closely related sympatric congener P. enesefae, the third group heard no frog sounds, and the fourth group heard only broadband white noise. At sexual maturity, we tested each female's preferences for conspecific complex vs simple calls and discrimination against calls of the sympatric congener. Female choices in all of these tests were consistent with those in previous studies of wild-caught and laboratory-reared specimens of this species. The acoustic rearing environments in this study did not alter the preferences of females for complex conspecific calls or the discrimination of females against the sympatric congener. This study supports the hypothesis that early experience does not alter the mate choice preferences of female P. pustulosus. [ABSTRACT FROM AUTHOR]

Abstract: Gonadal differentiation in frogs is affected by sex steroids and thyroid hormones (THs); however, the genes controlling differentiation and the molecular effects of THs in the gonad are not clear and have only been investigated in a few anuran species. In this study, we established developmental profiles and TH regulation of sex steroid- and TH-related genes in the gonad-mesonephros complex (GMC) of the túngara frog (Physalaemus pustulosus), and compared the results to our previous research in another tropical frog, Silurana tropicalis. The developmental profiles allowed us to identify three genes as markers of ovarian development. During metamorphosis, aromatase (cyp19), estrogen receptor α, and steroid 5α-reductase 1 (srd5alpha1) were higher in the GMC of putative and morphological females. Acute exposure to triiodothyronine (T3) decreased GMC expression of srd5alpha1 and cyp19, while increasing TH-related genes in premetamorphic tadpoles. The regulation of sex steroid-related genes differed significantly from our previous study in S. tropicalis. P. pustulosus and S. tropicalis share ecological, developmental, and reproductive characteristics; however, they are not closely related. These results along with our previous research in the tadpole brain support the hypothesis that evolutionary convergence is not important in understanding differences in the effects of TH on sex steroid-related genes in frogs. Finally, we propose that T3 induces male gonadal development but this can be achieved through different mechanisms depending on the species. [Copyright &y& Elsevier]

In amphibians, thyroid hormones (THs) are considered key regulators of brain remodeling during metamorphosis, while sex steroids (estrogens and androgens) control sexual differentiation and gonadal development. However, these two endocrine axes can interact during tadpole brain development. Previously, we demonstrated that THs affect sex steroid-related gene expression in the developing brain of Silurana tropicalis and Rana pipiens; however, the gene expression changes differed between species. We chose to study a third anuran species, Physalaemuspustulosus, to test new hypotheses about the role of THs in the regulation of brain gene expression. We first established developmental transcript profiles of TH- and sex steroid-related genes in the brain of P. pustulosus. Then, following the same protocols as in our previous studies, we investigated triiodothyronine (T3) regulation of brain transcripts in premetamorphic P.pustulosus and then compared the results with our previous two studies. In the case of TH-related genes, TH receptor beta (trbeta) and deiodinase type 3 (dio3), mRNA developmental profiles were similar in the three species and with respect to other species in the published literature. However, the profiles of TH receptor alpha (tralpha) and deiodinase type 2 (dio2) mRNA revealed differences between anuran species. Among the three anurans we have studied, the direction of the T3 regulation of TH-related genes was overall similar, but the magnitude of gene expression change differed depending on the rate of metamorphosis in a given species. For the sex steroid-related genes, each species exhibited similar developmental profiles but differed in their response to T3. In P. pustulosus, T3 reduced the expression of aromatase (cyp19) while increasing mRNA levels of androgen and estrogen receptors. These results are similar to previous research in R. pipiens but differ from data for S. tropicalis, for which we found an increase in androgen synthesis enzymes but no effect on cyp19. Together, we propose that T3 has the potential to induce the brain androgen system in anurans. This could be achieved by increasing androgen synthesis enzymes (S. tropicalis) or by decreasing estrogen synthesis (due to a decrease in cyp19 in P. pustulosus and R. pipiens). In conclusion, we demonstrated that mechanisms of hormone interactions differ between anuran species, but in all cases T3 appears to affect the balance of sex steroids in the brain, stimulating the androgen system. We have shown that the regulation of sex steroid-related genes by T3 is more similar among closely related species than species with similar reproductive and developmental characteristics. Copyright © 2011 S. Karger AG, Basel [ABSTRACT FROM AUTHOR]

In anurans, recognition of species-specific acoustic signals is essential to finding a mate. In many species, behavioral tests have elucidated which acoustic features contribute to species recognition, but the mechanisms by which the brain encodes these species-specific signal components are less well understood. The túngara frog produces a `whine' mating call that is characterized by a descending frequency sweep. However, much of the signal is unnecessary for recognition, as recognition behavior can be triggered by a descending two-tone step that mimics the frequency change in a portion of the whine. To identify the brain regions that contribute to species recognition in the túngara frog, we exposed females to a full-spectrum whine, a descending two-tone step that elicits recognition, the reversed two-tone step that does not elicit recognition, or no sound, and we measured expression of the neural activity-dependent gene egr-1 in the auditory brainstem and thalamus. We found that the behavioral relevance of the stimuli was the best predictor of egr-1 expression in the laminar nucleus of the torus semicircularis but not elsewhere. That is, the laminar nucleus responded more to the whine and the two-tone step that elicits recognition than to the reversed two-tone step. In contrast, in other brainstem and thalamic nuclei, whines induced egr-1 expression but tones did not. These data demonstrate that neural responses in the laminar nucleus correspond to behavioral responses of females and they suggest that the laminar nucleus may act as a feature detector for the descending frequencies characteristic of conspecific calls. [ABSTRACT FROM AUTHOR]

Factors that influence the strength of sexual selection within a population may have important consequences for the evolution and maintenance of reproductive isolation between populations. Here we examine the role of one such factor, predation risk, in influencing female permissiveness for heterospecific advertisement calls. Specifically, we tested the effect of simulated predation risk on mate choice in female túngara frogs, Physalaemus pustulosus, using variation in ambient light levels, travel time and the presence of auditory cues of local predators. Simulated predation risk increased female permissiveness for advertisement calls along an artificial gradient of calls intermediate between conspecifics and a congener, Physalaemus enesefae. Across the entire gradient, the presence of auditory cues of the predatory frog Leptodactylus pentadactylus in association with the conspecific call dramatically increased the likelihood of females choosing the intermediate call. In addition, higher ambient light levels and simulation of increased travel distances both increased the likelihood that females would choose intermediate calls over conspecific calls. These results suggest that, although mate choice may be important in causing reproductive isolation between allopatric populations, spatial or temporal variation in predation risk may strongly influence the expression of mate choice and thus the outcome of secondary contact. [Copyright &y& Elsevier]

Sexual selection is responsible for the evolution of costly elaborate male traits. When male displays are dynamic, display strategy is sensitive to contextual cues that alter the relative costs and benefits of producing each signal in a male’s repertoire. Because females often prefer more elaborate signals, males often respond to female presence by elaborating their display. When added elaboration increases assessment information or reproductive stimulation, females might benefit by extracting the maximum amount of signal elaboration from males. Thus, we expect that females could exaggerate their presence and cause males to produce even costlier and more attractive signals by exhibiting “elicitation” behaviors. We asked whether female túngara frogs elicit increased call complexity from prospective mates. In túngara frogs, adding complexity increases both attractiveness and predation risk. We found that females exhibit a repertoire of movements that function not in mate acquisition, per se, but in display manipulation, by eliciting increased complexity from calling males. The probability that males add complexity to their display increases when females produce these movements. Thus, females actively influence males to produce riskier signals. [ABSTRACT FROM PUBLISHER]

While the presence of predators can influence female mate choice, few studies have investigated how females respond to quantitative variation in predation risk. In addition, we know little of how females respond to multiple, independent cues of risk. In this study, we investigated the effects of simulated predation risk on mate choice in túngara frogs, Physalaemus pustulosus, using the advertisement calls of predatory frogs, variation in ambient light, and simulated distance. Females showed aversion to conspecific calls associated with the calls of predators, and females were significantly less likely to travel perceived longer distances while the calls of predatory frogs were broadcast. In both the laboratory and field, females chose among potential mates significantly faster under higher light levels. Female responses to acoustic cues of predation risk were significantly influenced by light level, but decisions about travel distances were not. These results demonstrate that female choice is strongly influenced by perceived predation risk and that females can simultaneously evaluate quantitative variation in different cues of predation risk. The changes in search behavior and mate evaluation we demonstrate indicate that predation plays a strong role in limiting signal evolution and possibly reproductive isolation. [ABSTRACT FROM AUTHOR]

In many species males vocally advertise for mates in choruses and these choruses serve as acoustic beacons to conspecific females as well as to eavesdropping predators and parasites. Chorusing will often cease in response to disturbances, such as the presence of predators. In some cases the cessation is so rapid and over such a large area that it seems improbable that males are all responding directly to the same local disturbance. Here, we demonstrate experimentally in Neotropical túngara frogs, Physalaemus pustulosus, that the cessation of calling by males spreads rapidly through the chorus. The cessation of chorusing in response to the cessation of playbacks of three calling males is more effective in inducing chorus cessation than is the cessation of one male calling. When three males are calling, the cessation of complex calls is more effective in inducing chorus cessation than simple calls. There is no main effect on whether the final call of the male is complete or is interrupted. We thus conclude that the sudden lack of signals-the 'sounds of silence'-becomes an alarm cue that can explain the rapid cessation of choruses that are common in many chorusing species. Abstract in Spanish is available at . [ABSTRACT FROM AUTHOR]

Abstract: The gonadal steroid hormone progesterone plays an important role across all vertebrates in mediating female reproductive physiology and behavior. Many effects of progesterone are mediated by a nuclear progesterone receptor (PR), which is crucial for integration of external signals and internal physiological cues in the brain to produce an appropriate behavioral output. The túngara frog, Physalaemus pustulosus, is an excellent model system for the study of mechanisms by which sensory signals, such as auditory communication, are processed within neural circuits where mate choice decisions are made. To establish a framework for studying the neural basis of mate choice and social behavior in this species, we first describe the cytoarchitecture of the brain using Nissl-stained sections. Then, in order to better understand where progesterone acts to regulate social decisions, we determined the distribution of PR protein throughout the brain of P. pustulosus by immunohistochemistry. We found PR immunoreactivity in key brain regions known to modulate the processing of auditory cues and social behavior in other vertebrates. Due to its widespread distribution, PR likely also plays important roles in non-limbic brain regions that mediate non-social information processing. Further, we have colocalized PR with tyrosine hydroxylase, providing a functional context for the role of progesterone in mediating motivation and motor behavior. Our results significantly extend our understanding of hormonal modulation in the anuran brain and support the important role of the nuclear progesterone receptor in modulating female mate choice and receptivity in amphibians and across vertebrates. [Copyright &y& Elsevier]

Disentangling the influence of multiple signal components on receivers and elucidating general processes influencing complex signal evolution are difficult tasks. In this study we test mate preferences of female squirrel treefrogs Hyla squirella and female túngara frogs Physalaemus pustulosus for similar combinations of acoustic and visual components of their multimodal courtship signals. In a two-choice playback experiment with squirrel treefrogs, the visual stimulus of a male model significantly increased the attractivness of a relatively unattractive slow call rate. A previous study demonstrated that faster call rates are more attractive to female squirrel treefrogs, and all else being equal, models of male frogs with large body stripes are more attractive. In a similar experiment with female túngara frogs, the visual stimulus of a robotic frog failed to increase the attractiveness of a relatively unattractive call. Females also showed no preference for the distinct stripe on the robot that males commonly bear on their throat. Thus, features of conspicuous signal components such as body stripes are not universally important and signal function is likely to differ even among species with similar ecologies and communication systems. Finally, we discuss the putative information content of anuran signals and suggest that the categorization of redundant versus multiple messages may not be sufficient as a general explanation for the evolution of multimodal signaling. Instead of relying on untested assumptions concerning the information content of signals, we discuss the value of initially collecting comparative empirical data sets related to receiver responses. [ABSTRACT FROM AUTHOR]

Facultative traits that have evolved under sexual selection, such as the acoustic ornaments present in the advertisement signals of male túngara frogs ( Physalaemus pustulosus), offer a unique opportunity to examine selection for trait exaggeration with a focus on individual differences amongst signalers. By contrast, many studies of mate choice use experimental designs that obscure the inter-individual variation amongst signalers available for selection to act on-through the use of 'typical' or average signals from the population. Here, we use dichotomous female phonotaxis choice tests to determine how the value of male call embellishment varies across 20 individual males frogs recorded from the wild-a sample which captures the acoustic diversity present in the population. We tested 20 females for each male call pair (i.e., 400 females). The results show widespread preference amongst females for ornamented calls ('whine-chucks') over simple calls ('whines'), yet also demonstrate substantial variation in the relative benefits for individual male frogs-some males enjoy appreciable benefits by using ornaments while others (30% of males in this study) do not. We also show that the relative amplitude of the chuck to the whine correlates positively with the value of call elaborations across these 20 males. Finally, by manipulating the relative amplitude of whines and chucks using both natural and synthetic calls, we demonstrate directly that this single call parameter is key to determining the relative value of call elaborations across males. [ABSTRACT FROM AUTHOR]

Search costs can have profound influences on female choice, causing females to become less choosy or sample less of the diversity of available mates. Predator foraging strategies, however, determine exactly how search time affects predator encounter rates. Ambush predators are more likely to be encountered by females traveling longer distances to evaluate males, but evaluation time is unlikely to influence encounter-rate with this type of predator. Actively searching predators, however, may be more likely to be encountered by females employing longer travel times and evaluation times. In this study, we examine the effects of perceived search costs on both temporal and spatial aspects of the search behavior of female túngara frogs, Physalaemus pustulosus. Females were collected from natural choruses and presented with conspecific calls at a distance of 50, 115, or 180 cm from their release point. Assays were conducted in either darkness or simulated full moon light levels. Longer starting distances caused longer choice latencies, but choice latency was considerably lowered under higher light conditions. Females spent considerably less time moving under higher light conditions; however, light levels did not affect path length. Females were more likely to leave the release point with more accurate orientation to the sound source under higher light conditions. We demonstrate that females can respond to perceived search costs by altering spatial and temporal aspects of female search behavior. The overall emphasis of females on reducing time spent moving and increasing movement speed indicates that predation by actively searching predators represents a stronger cost to females than ambush predators. [ABSTRACT FROM AUTHOR]

Dopamine is an evolutionarily ancient neurotransmitter that plays an essential role in mediating behavior. In vertebrates, dopamine is central to the mesolimbic reward system, a neural network concerned with the valuation of stimulus salience, and to the nigrostriatal motor system and hypothalamic nuclei involved in the regulation of locomotion and social behavior. In amphibians, dopaminergic neurons have been mapped out in several species, yet the distribution of dopaminoreceptive cells is unknown. The túngara frog, Physalaemus pustulosus, is an excellent model system for the study of neural mechanisms by which valuations of stimuli salience and social decisions are made, especially in the context of mate choice. In order to better understand where dopamine acts to regulate social decisions in this species, we have determined the distribution of putative dopaminergic cells (using tyrosine hydroxylase immunohistochemistry) and cells receptive to dopaminergic signaling (using DARPP-32 immunohistochemistry) throughout the brain of P. pustulosus. The distribution of dopaminergic cells was comparable to other anurans. DARPP-32 immunoreactivity was identified in key brain regions known to modulate social behavior in other vertebrates including the proposed anuran homologues of the mammalian amygdalar complex, nucleus accumbens, hippocampus, striatum, preoptic area, anterior hypothalamus, ventromedial hypothalamus, and ventral tegmental area/substantia nigra pars compacta. Due to its widespread distribution, DARPP-32 likely also plays many roles in non-limbic brain regions that mediate non-social information processing. These results significantly extend our understanding of the distribution of the dopaminergic system in the anuran brain and beyond. Copyright © 2010 S. Karger AG, Basel [ABSTRACT FROM AUTHOR]

The article presents a study which aims to examine how an alkaloid sequestration protect the green poison frog from predator attacks in Isla Taboga, Panama in 2010. Physalaemus pustulosus, a non-toxic frog, and Dendrobates auratus, an alkaloid-sequestering frog, were offered as prey to Sericopelma rubnitens, a theraphosid spider. The study reveals that an alkaloid sequestration did not prevent D. auratus from attacks, but did lessen the risk to be eaten. It shows that spiders are one of just a few recorded D. auratus predators in Taboga and might represent a strong inclusive force for Taboga's high toxicity.

In acoustically advertising anurans the male courtship call elicits species-typical responses from conspecifics - usually phonotactic approach and mate choice in gravid females and an evoked vocal response in adult males. Males in several species, however, are also known to perform phonotaxis, sometimes with the same acoustic preferences as females. Female túngara frogs are known to update their phonotactic approach as male advertisement signals change dynamically in attractiveness. Here we show that males also perform such temporal updating during phonotaxis in response to dynamic playbacks. While males exhibit slower phonotactic approaches than females, their responsiveness to dynamic changes in call complexity does not differ significantly compared to females. These results demonstrate that males are sensitive to the location of preferred call types on a moment-to-moment basis and suggest that similarities between male and female sexual behaviour in anurans might often be overlooked. We suggest that anuran phonotaxis is more widespread and serves different functions in reproductive females and males. Lastly, these temporal updating results suggest that male frogs are highly selective about site selection in a chorus. [ABSTRACT FROM AUTHOR]

Males and females can differ both in the social behaviors they perform and in the contexts in which they engage in these behaviors. One possible mechanism of sex differences in behavior is a sexual dimorphism in the relay of sensory information to motor areas, but no studies have examined the role of such a relay in vertebrate sexually dimorphic behaviors. We used egr-1 expression as a marker of neural activation in frogs exposed to conspecific and heterospecific acoustic signals to compare activation patterns throughout the brains of males and females. We determined how the sexes differ in the transformation of social signals into motor responses in the context of social communication. We examined the relationships between egr-1 mRNA levels in the auditory midbrain and forebrain areas, as well as how forebrain expression related to the behavioral responses of the animals. Forebrain network activation patterns and forebrain-behavior relationships were similar in males and females. By contrast, we found a sex difference in the relationship between midbrain and forebrain activation; midbrain auditory responses predicted forebrain responses in females but not in males. This sex difference suggests that sensory inputs differentially regulate motor systems underlying social behaviors in males and females. This sensorimotor transformation may be a common locus for generating sex differences in behavior. [ABSTRACT FROM AUTHOR]

Background: During mate choice, individuals must classify potential mates according to species identity and relative attractiveness. In many species, females do so by evaluating variation in the signals produced by males. Male túngara frogs (Physalaemus pustulosus) can produce single note calls (whines) and multi-note calls (whine-chucks). While the whine alone is sufficient for species recognition, females greatly prefer the whine-chuck when given a choice. Methodology/Principal Findings: To better understand how the brain responds to variation in male mating signals, we mapped neural activity patterns evoked by interspecific and intraspecific variation in mating calls in túngara frogs by measuring expression of egr-1. We predicted that egr-1 responses to conspecific calls would identify brain regions that are potentially important for species recognition and that at least some of those brain regions would vary in their egr-1 responses to mating calls that vary in attractiveness. We measured egr-1 in the auditory brainstem and its forebrain targets and found that conspecific whine-chucks elicited greater egr-1 expression than heterospecific whines in all but three regions. We found no evidence that preferred whine-chuck calls elicited greater egr-1 expression than conspecific whines in any of eleven brain regions examined, in contrast to predictions that mating preferences in túngara frogs emerge from greater responses in the auditory system. Conclusions: Although selectivity for species-specific signals is apparent throughout the túngara frog brain, further studies are necessary to elucidate how neural activity patterns vary with the attractiveness of conspecific mating calls. [ABSTRACT FROM AUTHOR]