Mate sampling by female barking treefrogs (Hyla gratiosa)

Despite intense interest in mate choice, relatively little isknown about how individuals sample prospective mates. Indeed,a key issue is whether females sample males or simply matewith the first male encountered. We investigated mate samplingby female barking treefrogs (Hyla gratiosa). Females choosingmates in natural choruses did not move between males but insteadmated with the first male they approached closely. Most femalesmated with the male closest to them at the start of their mate-choiceprocess, and females were more likely to mate with the closest male when the distance to other males was large. These observationsare consistent with the hypothesis that females do not samplepotential mates but instead mate with the first male they distinguishfrom the rest of the chorus. To test this initial detectionhypothesis, we conducted a playback experiment in which weoffered females a choice between two calls, one of which was detectable above the background chorus sound at the female'srelease point, and one of which became detectable only as femalesmoved toward the initially detectable call. Females did notprefer the initially detectable call, thus ruling out the initialdetection hypothesis and implicating sampling of potentialmates by females. Based on the behavior of females in natural choruses, we hypothesize that females approach the chorus, moveto locations where they are able to detect the calls of severalmales simultaneously, and choose a mate from among these malesat some distance from the males. Such simultaneous samplingmay be common in lekking and chorusing species, which havebeen the subjects of many studies of sexual selection.     

Hormone-induced vocal behavior and midbrain auditory sensitivity in the green treefrog,Hyla cinerea

Summary Twenty four castrated male, 6 intact male, and 11 intact female Hyla cinerea were injected subcutaneously with 25 g arginine-vasotocin (AVT) and induced to call 1 h later in response to the playback of a conspecific mating call. Eighteen castrated males and 8 intact females were implanted 5 mg androgen pellets for 3 weeks prior to the neuropeptide injection. Among castrated males, 6/9 testosterone (T) implanted, 4/9 dihydrotestosterone (DHT) implanted and 2/6 non implanted individuals produced calls after being administered AVT. 5/6 intact non implanted males and 6/8 T intact implanted females also called, and 3 intact non implanted females remained silent after the injection. Evoked calls had a mid-frequency spectral peak at about 1900 Hz which is absent in field-recorded mating calls of this species. Calls of implanted females and castrated non implanted males were shorter than those of castrated implanted and intact non implanted males. Audiograms measured before hormone implants showed dips of enhanced sensitivity at about 0.5, 0.9 and 3.0 kHz in males and females. After AVT injection, thresholds at frequencies within the 0.7–1.5 kHz range were increased in castrated males. Such reduction in sensitivity points to an inhibition of the auditory system during hormone induced vocal activation.Abbreviations AVT arginine-vasotocin - DHT dihydrotestosterone - T testosterone - TS torus semicircularis     

Frequency selectivity of hearing in the green treefrog,Hyla cinerea

Frequency selectivity of hearing was measured in the green treefrog, Hyla cinerea. A psychophysical technique based on reflex modification was used to obtain masked threshold estimates for pure tones (300-5,400 Hz) presented against two levels of broadband masking noise. A pure tone (S-1) presented 200 ms prior to a reflex-eliciting stimulus (S-2) inhibited the motor reflex response to S-2. The magnitude of this reflex modification effect varied systematically with the sound pressure level (SPL) of S-1, and threshold was defined as the SPL of S-1 at which the reflex modification effect disappeared. Masked thresholds were used to calculate critical ratios, an index of the auditory system's frequency selectivity. The frequency selectivity of the treefrog's hearing is greatest and critical ratios are lowest (22-24 dB) at about 900 and 3,000 Hz, the two spectral regions dominant in the male treefrog's species-specific advertisement call. These results suggest that the treefrog's auditory system may be specialized to reject noise at biologically-relevant frequencies. As in other vertebrates, critical ratios remain constant when background noise level is varied; however, the shape of the treefrog's critical ratio function across frequencies differs from the typical vertebrate function that increases with increasing frequency at a slope of about 3 dB/octave. Instead, the treefrog's critical ratio function resembles its pure tone audiogram. Although the shape of the treefrog's critical ratio function is atypical, the critical ratio values themselves are comparable to those of many other vertebrates in the same frequency range. Critical ratio values here measured behaviorally do not match critical ratio values previously measured physiologically in single eighth nerve fibers.(ABSTRACT TRUNCATED AT 250 WORDS)     

Selectivity for harmonic structure in complex sounds by the green treefrog (Hyla cinerea)

1. A psychophysical technique based on reflex modification was used to study the detection of two-tone complexes in background noise by the green treefrog (Hyla cinerea). Three different two-tone complexes were synthesized and presented to measure detection thresholds--a harmonic complex of 900 + 3000 Hz (periodicity of 300 Hz, mimicking the structure of the natural advertisement call); an inharmonic complex of 830 + 3100 Hz; and a second harmonic complex of 828 + 2760 Hz (periodicity of 276 Hz). 2. Masked thresholds and 'critical ratios' (signal-to-noise ratios at threshold) were lowest for the two harmonic complexes (900 + 3000 Hz, mean 'critical ratio' of 16 dB; 828 + 2760 Hz, mean 'critical ratio' of 14 dB). For the inharmonic complex, for which there is no stable first-harmonic periodicity, the mean 'critical ratio' was 24 dB. These data suggest that the green treefrog is sensitive to the harmonic structure of complex sounds as a specific acoustic feature. 3. Because of the unique structure of the treefrog's inner ear, the heightened behavioral sensitivity to harmonic complexes must be due to processing in the central, rather than peripheral, auditory system.     

Social signals influence hormones independently of calling behavior in the treefrog (Hyla cinerea)

Social signals play an important role in regulating hormone-behavior relationships. In anurans (frogs and toads), acoustic signals are an essential aspect of reproductive behavior; however, the physiological consequences of receiving social signals has remained largely undescribed. Each night for 5, 10, or 20 days, we presented acoustically isolated male treefrogs with a conspecific mating chorus, an array of tones, or no sound. We recorded calling rate of individuals throughout the experiment and collected blood before and after treatment. Days of stimulus exposure had no effect on any dependent measure. Acoustic treatment influenced steroid levels; testosterone, dihydrotestosterone, and corticosterone increased only in the group exposed to the chorus. Chorus-exposed males also showed an increase in stimulus-evoked calling. We found no correlation between androgens and calling within each treatment group. In addition, noncallers in the chorus group had higher levels of androgens than males in the tone or no sound groups. Further, chorus-exposed males with zero, low, or high rate of calling had similar levels of androgens. These data indicate that social signals increase circulating androgens independently of calling behavior. Elevated corticosterone associated with chorus reception did not inhibit calling behavior, and corticosterone showed no correlation with androgen levels.     

The separate and combined effects of harmonic structure, phase, and FM on female preferences in the barking treefrog (Hyla gratiosa)

In natural advertisement calls of the barking treefrog, Hyla gratiosa, a small amount of incoherent frequency modulation (FM) is present. Incoherency in the FM of a call creates inharmonicity and phase changes between its frequency components. In this study, the combined and separate effects of the harmonic structure, phase spectrum, and FM of an advertisement call on female choice were tested. The harmonic structure of a call can have a direct effect on female preference; females showed a significant preference for static-inharmonic calls over static-harmonic calls. Neither differences in phase or FM alone conferred a preference in two choice tests. However, when FM is present in both calls it does influence female preference for harmonic structure -namely harmonic calls become preferable to inharmonic calls. This reversal of female preference for inharmonicity in a call by the presence of FM suggests that call parameters may interact, and thereby effect mate choice.Abbreviations AP amphibian papilla - BP basilar papilla - FM frequency modulation - PM phase modulation - HS harmonic structure - GB Gaussian Band     

Field and laboratory studies of the susceptibility of the green treefrog (Hyla cinerea) to Batrachochytrium dendrobatidis infection

Amphibians worldwide are experiencing devastating declines, some of which are due to the amphibian chytrid fungus (Batrachochytrium dendrobatidis, Bd). Populations in the southeastern United States, however, have not been noticeably affected by the pathogen. The green treefrog (Hyla cinerea) is abundant and widespread in the southeastern United States, but has not been documented to harbor Bd infection. This study examined the susceptibility of H. cinerea to two strains of Bd in the lab and the prevalence of infection in wild populations of this species in southeastern Louisiana. Although we were able to infect H. cinerea with Bd in the lab, we did not observe any clinical signs of chytridiomycosis. Furthermore, infection by Bd does not appear to negatively affect body condition or growth rate of post-metamorphic individuals. We found no evidence of infection in surveys of wild H. cinerea. Our results suggest that H. cinerea is not susceptible to chytridiomycosis post-metamorphosis and probably is not an important carrier of the fungal pathogen Bd in the southeastern United States, although susceptibility at the larval stage remains unknown.     

Acoustic modulation of neural activity in the preoptic area and ventral hypothalamus of the green treefrog (Hyla cinerea)

Summary Responses of neurons in the preoptic area and ventral hypothalamus to conspecific mating calls or white noise bursts were examined in male green treefrogs (Hyla cinerea) during different seasons. In the winter, 34.3% of preoptic neurons and 46.7% of ventral hypothalamic cells demonstrated significant changes in activity level during presentation of a conspecific mating call. In contrast, only 13.3% of preoptic units and 16.7% of ventral hypothalamic cells responded to the white noise. The percentage of preoptic and hypothalamic units responding to the advertisement call did not differ significantly during the summer breeding season. Type I units exhibited a dramatic increase in activity during acoustic stimulation followed by a rapid return to baseline activity levels after stimulus offset. Type II cells showed a robust activity increase during stimulation, but maintained an intermediate activity level after stimulus offset. In the preoptic area, a third response type exhibited suppressed activity during acoustic stimulation. Although seasonal condition did not alter the percentage of acoustically responsive units within either nucleus, the proportion of Type I units in the ventral hypothalamus was greatest during the summer.Abbreviations MC mating call - NS no stimulus - POA preoptic area - VH ventral hypothalamus - WN white noise     

Proximate developmental causes of limb length variation between Hyla cinerea and Hyla gratiosa (Anura: Hylidae)

Hyla cinerea and H. gratiosa are closely related treefrogs that differ at metamorphosis in the length of the hind limb relative to snout vent length. A developmental series of larvae of the two species raised under uniform laboratory conditions reveals that H. cinerea have proportionately longer legs at metamorphosis than H. gratiosa for the following reasons: 1) Hyla cinerea initiate limb growth at a smaller body size than H. gratiosa, 2) H. cinerea have a faster rate of limb growth, and 3) body growth rates in H. cinerea and H. gratiosa are nearly identical until just before metamorphosis.     

The cause of correlations between nightly numbers of male and female barking treefrogs (Hyla gratiosa) attending choruses

The number of males displaying in a lek or chorus each day isoften positively correlated with the number of females visitingor mating in the aggregation. I tested hypotheses that mightexplain such correlations in a study of the barking treefrog(Hyla gratiosa). Experimental reduction of the number of callingmales did not reduce female visitation, ruling out the hypothesisthat such correlations are owing to female preference for, orpassive attraction to, larger choruses. Separate regressionof the numbers of males and females on 13 environmental variablesexplained 45–74% of the variance in the nightly numbersof males and females. Partial regression coefficients for mostof the 13 variables were not significantly different betweenthe sexes, and the relative importance of variables in explainingvariation in the numbers of individuals was similar for bothsexes. These results support the hypothesis that positive correlationsbetween nightly numbers of males and females are owing to asimilar response by both sexes to the same environmental variables.Thus, it appears that the intense sounds emanating from chorusesof H. gratiosa do not function in long-range communication andmay instead be an epiphenomenon of intense, short-range vocalcompetition for females.     

Reproductive character displacement in the acoustic communication system of green tree frogs (Hyla cinerea)

Abstract Interactions between species can affect the evolution of their sexual signals, receiver selectivity, or both. One commonly expected outcome is reproductive character displacement, whereby adverse consequences of mismating select for greater differentiation of communication systems in areas of sympatry than in areas of allopatry. We found evidence of reproductive character displacement in the acoustic communication system of green tree frogs ( Hyla cinerea ). The strength of female preferences for the spectral properties of calls that distinguish conspecific calls from those of a closely related congener, H. gratiosa , was greater in areas of sympatry with H. gratiosa than in areas of allopatry. We also found subtle differences in advertisement calls and in the heights of male calling perches when we restricted our comparisons to localities in which H. gratiosa was also breeding (syntopy) with localities where this species was absent. Hyla cinerea and H. gratiosa show only weak genetic incompatibility, but the calls representative of interspecific hybrids were unattractive to females of both parental species. Hybrids might also be at an ecological disadvantage because of different habitat preferences of the two taxa. Thus, selection against production of less fit or less attractive hybrid or backcross offspring are probably the main causes responsible for the differences documented in this paper.     

Success of intracoelomic radiotransmitter implantation in the treefrog (Hyla versicolor)

Concern about the effects of habitat modification on the natural movements and behavior of amphibians has led researchers to devise several methods to follow individual animals as they migrate among seasonal resources; one such method is the surgical insertion of radiotransmitters. Intracoelomic implantation has come into general use despite the relative lack of specific procedural guidelines and information about the effects of transmitter insertion and removal. The author outlines the surgical implantation procedures used on a small amphibian (gray treefrog) and provides insight into the effects of multiple major surgical invasions on the long-term survival of amphibians. Multiple survival surgeries were successful in this study and may be an alternative to euthanasia in some instances.