Temporal and Geographic Variation in the Advertisement Call of the Booroolong Frog (Litoria booroolongensis: Anura: Hylidae)

The mechanisms that underlie sexual selection rely upon within‐ and among‐individual variability in the targeted traits. In this study, we examined variation in the advertisement call of the booroolong frog (Litoria booroolongensis) at several different levels: between populations, between breeding seasons in the same population, among males within a population, within males between nights and within males in a single calling bout. The call of L. booroolongensis has multiple notes with a pulsed structure. We detected considerable variation in advertisement call structure between breeding seasons and between populations. The measured call properties ranged from static to dynamic; however, most properties were intermediate between the criteria that have been traditionally used to define call traits as static or dynamic (≤5 and ≥12% respectively). We compared actual and relative repeatabilities and found that the temporal call properties associated with the structure of the note had the highest values, suggesting that these characters in particular may respond to selection. We argue that relative repeatabilities are a particularly useful measure of the potential for evolutionary response to selection as they account for an individual's relative performance during the period of assessment in an ever‐changing breeding arena.     

The Advertisement Call of the European Treefrogs (Hyla arborea): A Multilevel Study of Variation

In this study, we analysed the patterns of variation of the European treefrog's advertisement call at four levels of organization: within individuals, within populations, among populations of the same species, and among different species of the same clade. At the within-individual level, call acoustic properties are distinguished into static and dynamic properties. At the within-population level, two sources of call variation were analysed: temperature and body size. Temperature affects both temporal and spectral properties of the call. Body size mostly affects spectral properties. At the between-population level, calls do not show significant differences with respect to temporal properties, but they do differ in two spectral (and stereotyped) properties: the fundamental frequency and the difference in amplitude between dominant and fundamental frequencies. Finally, at the between-species level, call differences are much more conspicuous: they involve both spectral and temporal and both static and dynamic properties. At all four levels, body size is associated with call variation, explaining 11% of the differences among populations of the same species and 73% of the differences among species of the same clade. On the basis of these results, we hypothesize that patterns of variation of call acoustic properties, their constraints, and their biological functions are intrinsically associated. We discuss the role that such an association might play in the evolution of acoustic signals.     

Individual Variation in Advertisement Calls of Territorial Male Green Frogs, Rana clamitans: Implications for Individual Discrimination

Individuals of many territorial species discriminate between familiar territorial neighbors and unfamiliar strangers based on individual differences in acoustic signals. Many anuran amphibians are territorial and use primarily acoustic communication during social interactions, but evidence for acoustically mediated individual discrimination is available only for one species. As a first step in research designed to investigate individual discrimination in a second species of territorial frog, we examined patterns of within-male and among-male variability in 198 advertisement calls of 20 male green frogs, Rana clamitans . The aim was to determine which acoustic properties could be used by males to recognize their territorial neighbors and to estimate limits of reliability afforded by these properties for identifying different neighbors. All of the call properties that we examined exhibited significant inter-individual variation. Discriminant function analyses assigned between 52% and 100% of calls to the correct individual, depending on sample size and the call properties included in the model. This suggests that there is sufficient among-male variability to statistically identify individuals by their advertisement calls. The call properties of fundamental frequency and dominant frequency contributed the most towards discrimination among individuals. Based on their natural history and behavior and the results reported here, we suggest that male green frogs likely discriminate between strangers and adjacently territorial neighbors based on individual variation in advertisement calls.     

Differential Evolution of Advertisement Call Traits in Dart-Poison Frogs (Anura: Dendrobatidae)

The ability to recognise and discriminate between heterospecific and conspecific individuals plays an essential role in mate choice, reproductive isolation and thus species diversification. Many animals discriminate based on advertisement calls, whose evolution may be driven by a variety of forces such as natural selection, sexual selection or stochastic processes. The relative importance of stochastic processes acting on a given trait is usually correlated with its phylogenetic signal. Mate-recognition signals are complex traits composed of multiple features that could potentially respond independently to evolutionary forces. The advertisement call of anurans is used in species recognition and mate choice. In this study, we estimate the phylogenetic signal for body size and a suite of traits describing the male advertisement call from dart-poison frogs (Anura: Dendrobatidae). We found a surprisingly high phylogenetic signal for all call traits. In addition, call traits varied in their degree of phylogenetic signal, suggesting that evolutionary forces have been acting differently on different traits. Pulse duration showed the strongest phylogenetic signal. Peak frequency and body size were correlated and presented high phylogenetic signal indicating that the evolution of one trait may be driving or constraining the other. Since most variation in call traits can be explained by the phylogenetic history of the species, we cannot reject the hypothesis that stochastic processes account for significant evolutionary divergence in frog calls.     

No Evidence for the Compensation Hypothesis in the Swelled Vent Frog(Feirana quadranus)

The compensation hypothesis predicts that if the left testis is defective e.g. due to developmental stress,the increased right testis serves a compensatory role, and thereby displaying testes asymmetry which can be a reliable indicator of male body condition. Here, to test the prediction of the compensation hypothesis, we analyzed difference in size between left testis and right testis and the relationship between testes asymmetry and male body condition in the swelled vent frog(Feirana quadranus).We found that the left testis was larger than right testis,displaying a significant directional asymmetry in testes size. Although testes mass was correlated with body condition, testes asymmetry was not correlated with body condition, which cannot provide evidence that the right testis had a compensatory function. Our findings suggest no evidence for the compensation hypothesis in this species due to lacking the compensatory function in right testis.     

Ring Clinal Variation in Morphology of the Green Odorous Frog(Odorrana margaretae)

The green odorous f rog(Odorrana margaretae) has an interesting ring-shaped divergence pattern around the Sichuan Basin and documenting its morphological variations is essential in understanding its evolutionary history. Using curvilinear models,we detected significant geographical clinal variations in morphological traits, particularly sizes, of female O. margaretae. Males had significantly smaller sizes than females, and also had smaller variation ranges than females. One major trend of morphological variations was clinal: populations f rom the west tended to have a larger size with wider head and longer posterior limbs than populations from the east.Species history, with an early extended isolation and two subsequent secondary contacts, may explain most of the geographical clinal variations of O. margaretae.Bioclimatic factors may also contribute in explaining the variance of morphology.     

Pillar Function in the Fiddler Crab Uca beebei (II): Competitive Courtship Signaling

Male Uca beebei court and attract females into burrows they defend on muddy sand flats in the intertidal zone on the Pacific coast of the tropical Americas. Mating, oviposition and incubation (breeding) occur underground in males' burrows. Some courting males build mud pillars (2 cm high) at the entrance of their burrow. The purpose of this field study was to assess the role of pillars in competitive courtship signaling among males. I studied the effect of pillars on female behavior by recording the responses of wandering females to courtship from males resident at burrows with and without pillars. I also caught females, released them individually in a circular arena with an equal number of empty burrows with and without pillars around its circumference, and chased the females with a simulated avian predator. Females moved to burrows of both types more often when they were courted (82%) than when they were chased (67%). Receptive females were attracted to the burrows of the males that courted them significantly more often (97%) when these burrows had pillars than when they lacked pillars (66%). However, once females entered males' burrows they were equally likely to remain, mate and breed in both types of burrows. Females also more often moved to burrows with pillars (66%) than to burrows without pillars when they ran from the simulated predator. Both male courtship displays and pillars probably provide cues females use to locate males' burrows. The visual similarity between pillars and a display courting males give immediately before they enter their burrows suggests that pillars are icons of the display. The effect of pillars on female behavior, the timing of pillar building relative to when females choose mates, and contrasts in the behavior of males that do and those that do not build pillars suggest that pillar building has evolved due to competition among males to attract females into their burrows.     

Calls of the Large Odorous Frog(Odorrana graminea)Show Nonlinear and Individual Characteristics

Male large odorous frogs(Odorrana graminea)have been shown to have a rich vocal repertoire and their calls contain harmonics that extend into the ultrasonic range,much like calls of male O.tormota,a sympatric species.Whereas vocalizations of male O.tormota exhibit nonlinear characteristics and contain individual signatures,it is unclear whether or not those of O.graminea do.We carried out an extensive study of the vocalizations of male O.graminea with the goal of investigating whether they:(1)contained nonlinear phenomena(NLP),(2)carried individual signatures.We recorded and analyzed 1770 vocal calls from nine males.Their calls were classifiable into two distinct categories(one-note calls and multi-note calls).Of these,one-note complex/steep-frequency modulation call was the most abundant call-type.These calls showed diverse frequency modulation patterns,and abundant presence of NLP.Various basic spectral and temporal pa ra meters of such calls were individually distinct.Kruskal-Wallis Analysis of Rank test showed that the calls from individual males differed significantly.Subsequent discriminant function analysis(DFA)showed that 54.7%of the calls could be correctly assigned to individual.These results suggested that male O.graminea might be able to discriminate neighboring frogs from strangers acoustically.However,further research is necessary to demonstrate this capacity behaviorally.     

Altitudinal Variation in Digestive Tract Length in Yunnan Pond Frog （Pelophylax pleuraden）

The digestive tract plays an important role in digestion and the acquisition of food energy. Understanding the impact of abiotic environments on digestive tract morphology is especially important for evolution of digestive tract across different environments. Here, we investigated altitudinal variation in digestive tract length in the Yunnan Pond Frog （Pelophylaxpleuraden） across five populations ranging from 1413 m to 1935 m a.s.1, in Ningnan County, Sichuan province in western China. Frogs were collected during the breeding season, from 1-5 June 2012. Our results revealed that females had longer digestive tract and relative digestive tract （i.e. digestive tract length / body size） lengths in com- parison to males, on average, but the differences between them decreased with increasing altitude. Digestive tract and relative digestive tract lengths increased with increasing altitude suggesting that a higher proportion of indigestible ma- terials may be consumed at high-altitude sites and result in a relative increase in digestive tract dimensions.     

Functional Significance of Labellum Pattern Variation in a Sexually Deceptive Orchid (Ophrys heldreichii): Evidence of Individual Signature Learning Effects

Mimicking female insects to attract male pollinators is an important strategy in sexually deceptive orchids of the genus Ophrys, and some species possess flowers with conspicuous labellum patterns. The function of the variation of the patterns remains unresolved, with suggestions that these enhance pollinator communication. We investigated the possible function of the labellum pattern in Ophrys heldreichii, an orchid species in which the conspicuous and complex labellum pattern contrasts with a dark background. The orchid is pollinated exclusively by males of the solitary bee, Eucera berlandi. Comparisons of labellum patterns revealed that patterns within inflorescences are more similar than those of other conspecific plants. Field observations showed that the males approach at a great speed and directly land on flowers, but after an unsuccessful copulation attempt, bees hover close and visually scan the labellum pattern for up to a minute. Learning experiments conducted with honeybees as an accessible model of bee vision demonstrated that labellum patterns of different plants can be reliably learnt; in contrast, patterns of flowers from the same inflorescence could not be discriminated. These results support the hypothesis that variable labellum patterns in O. heldreichii are involved in flower-pollinator communication which would likely help these plants to avoid geitonogamy.     

The Complex Acoustic Design of an Advertisement Call in Male Mouse Lemurs (Microcebus murinus,Prosimii, Primates) and Sources of its Variation

During the breeding season sexually active male mouse lemurs produce a trill call used for advertising. The trill call displays a variable, high pitched, complex frequency and amplitude modulated acoustical structure and is given with a high sound intensity. The different sources of its variation in structure and usage have to be determined to understand its complex acoustical design and function. We investigated the vocal morphology of the trill call of males with known age and weight as well as the behavioral context, in order to evaluate sources of its variation. Cluster and discriminant analyses revealed that the structure of trill calls showed a high degree of individual stereotypy in contrast to a high interindividual variability. A set of three distinct variables (fundamental frequency of call onset, bandwidth of 3rd frequency modulation and end frequency of 7th modulation) was sufficient to identify individual senders with almost 90% probability. The structure of the trill call of adult males was independent of age, weight and context and was stable over at least one breeding season. Its individually distinct call structure and use in relation to social status provide the potential for individual recognition and for assessing the “quality” of a male.     

Pillar Function in the Fiddler Crab Uca beebei (I): Effects on Male Spacing and Aggression

Males of 14 species of fiddler crabs (genus Uca) are known to build structures out of mud or sand at the entrances to burrows they court from and defend. A study of spacing, space use and aggression between courting male Uca musica (Zucker 1974, 1981) suggested that the hoods males build reduce territorial overlap and rates of aggression between neighboring males. Thus, each male may have more time to court females during limited lunar, diurnal and tidal mating periods. I studied the courtship and aggressive behavior of male Uca beebei in the field to determine if the pillars males of this species build affect male behavior as do the hoods of U. musica. U. beebei occurs sympatrically with U. musica on the Pacific coast of Panama and is broadly similar in its ecology and mating behavior. Unlike the hoods of U. musica, pillars did not focus a male's activity space away from its closest neighbor nor did they reduce either overlap with neighbors' activity spaces or rates of aggressive interaction among neighbors. Pillar builders courted more but also fought their neighbors more than did males that did not build pillars. The pillars of U. beebei and the hoods of U. musica affect male behavior differently and probably have different functions.     

Individual Variation in Trophic Egg Production: Evidence for Maternal Manipulation in Response to Resource and Competition Levels

Females of the subsocial shield bug Parastrachia japonensis (Heteroptera: Parastrachiidae) incorporate trophic eggs (nutritive eggs) into their egg mass. Considerable variation occurs among females in trophic egg number and the proportion of an egg mass that is composed of trophic eggs. Because trophic eggs are essential to the development and survival of young, this variation could significantly impact female fitness. We tested the hypothesis that trophic egg abundance is induced by maternal phenotype (weight, body size) and resource exposure. We predicted that resource limitations would cause females to produce fewer fertile eggs and more trophic eggs and that larger and heavier females would produce more of each egg type. Females ovipositing early in the season are exposed to different resource conditions than those that oviposit late. Thus, we compared egg production patterns between these two groups and several other factors related to nesting. No correlation was seen between body size and trophic egg abundance, or, indeed, egg production, overall; however, heavier females produced heavier egg masses. Counter to our prediction, late females, which had greater access to food, produced significantly more total eggs, fewer fertile eggs, and more trophic eggs than early females. A binomial generalized linear model analysis indicated that the factors most correlated with the percentage of an egg mass destined to become trophic eggs were resource abundance, resulting from early or late oviposition, and distance of the nest from the host tree, with closer females producing more trophic eggs. The findings support our hypothesis that resource availability and, to a lesser extent, maternal phenotype affect trophic egg abundance.     

The Consequences for Communication of Call Overlap in the Tungara Frog,a Neotropical Anuran with a Frequency-modulated Call

We presented male tungara frogs, Physalaemus pustulosus, with call-triggered computer-synthesized whines that either did or did not overlap the frogs' own frequency-modulated whines in time. When the stimulus was nonoverlapping, males added a high proportion of chuck notes to their calls. When the stimulus was overlapping, males responded with either no or only small increases in chucking. This suggests they have difficulty detecting whines while vocalizing. When female tungara frogs were given a choice between alternating whines and out-of-phase overlapping whines they did not discriminate. The result with males was similar to that previously obtained with species of frogs that have amplitude-modulated advertisement calls (Schwartz 1987a), the result with females was different. We discuss our findings in relation to peripheral processing of spectral and temporal information in the anuran auditory system.     

Genetic Variation in Captive Koalas (Phascolarctos cinereus): Parentage Determination and Individual Identification

Highly repeatable randomly amplified polymorphicDNA (RAPD) markers were developed for parentage studiesin the koala (Phascolarctos cinereus). Of the 25 RAPDprimers screened, 5 (20.0%) produced 32 repeatable polymorphic RAPD bands (average/primer = 6.4± 4.2). A high level of polymorphism was observedfor each group of koalas (Featherdale, 71.9%; Lone Pine,84.4%). All 25 koalas could be uniquely identified using either RAPD or microsatellite markers. Of the32 RAPD markers generated in koalas, 25 were informativefor parentage analyses. These RAPD markers successfullydetermined both parents to three offspring and a male parent to a fourth offspring.Paternity analysis (where the female parent is known)succeeded in assigning the correct male parent to sevenoffspring. Our RAPD–PCR method generatesinformative genetic markers that are useful for parentagedetermination and individual identification of captivekoalas. This would provide genetic analysis to zoos andwildlife parks as a low-cost alternative to the more expensive microsatellite markers.     

Sound or Silence: Call Recognition in the Temporal Domain by the Frog Allobates femoralis

Acoustic communication often mediates agonistic interactions in territorial species. Because both the reaction to potential intruders and the lack thereof are costly, mechanisms that allow recognition of conspecific signals should be evident in intrasexual communication systems. While the spectral domain of the recognition space of the frog Allobates femoralis appears asymmetrically shaped in a way that reduces masking interference by the often syntopic frog Ameerega trivittata, frequency alone does not appear to account for the correct identification of conspecific intruders (Amézquita et al., Animal Behaviour, 70, 2005: 1377). As signal recognition may rely on a combination of spectral and temporal parameters of the signal, we test here the subsequent prediction that the recognition space should be asymmetrical in the temporal domain as well. We conducted playback experiments with 80 synthetic calls on 30 males and modeled all‐or‐none responses to define unidimensional functions of the recognition space for two call parameters: note duration and internote interval. For both parameters, male maximal response matched very well the average values of the conspecific signal and decreased with concomitant deviations from these values. While the response curves exceeded the range of signal variation and were not asymmetrical for either call parameter, they differed in breadth. The highest male permissiveness to variation in internote interval, as evidenced by a broader response curve, coincided with the lower probability of between‐species overlap in this signal parameter. Together with previous studies, our data suggest that a combination of spectral and temporal parameters of the advertisement call is necessary for recognition of calling intruders in A. femoralis. Our results emphasize the importance of multidimensional approaches in understanding signal recognition mechanisms in acoustically complex environments.     

Diversity of the Vocal Signals of Concave-Eared Torrent Frogs (Odorrana tormota): Evidence for Individual Signatures

Male concave-eared torrent frogs ( Odorrana tormota ) have an unusually large call repertoire and have been shown to communicate ultrasonically. We investigated the individual specificity of male advertisement calls in order to explore the acoustic bases of individual recognition, which was demonstrated in an accompanying study. Vocalizations of 15 marked males were recorded in the field. A quantitative analysis of the signals revealed eight basic call-types. Two of them (the single- and multi-note long-calls) were investigated in more detail. Long-calls were characterized by pronounced and varying frequency modulation patterns, and abundant occurrence of nonlinear phenomena (NLP), i.e., frequency jumps, subharmonics, biphonations and deterministic chaos. The occurrence of NLP was predictable from the contour of the fundamental frequency in the harmonic segment preceding the onset of the NLP, and this prediction showed individual-specific patterns. Fifteen acoustic variables of the long calls were measured, all of which were significantly different among individuals, except biphonic segment duration. Discriminant function analysis (DFA) showed that 54.6% of the calls could be correctly assigned to individual frogs. The correct classification was above chance level, suggesting that individual specificity of calls underlie the ability of males to behaviorally discriminate the vocal signals of their neighbors from those of strangers, a remarkable feat for a frog species with a diverse vocal repertoire. The DFA classification results were lower than those for other anurans, however. We hypothesize that there is a tradeoff between an increase in the fundamental frequency of vocalizations to avoid masking by low-frequency ambient background noise, and a decrease in individual-specific vocal tract information extractable from the signal.     

Call Variation in Tufted Titmice (Baeolophus bicolor) in a Distress Context: Potential Sex Differences

Tufted titmice, Baeolophus bicolor, produce calls in the contexts related to threat and approach of, and capture by, a predator. In titmice, these calls transition from the chick‐a‐dee call, used in a wide range of social contexts, to ‘distress’ calls that are produced by birds when captured and held by a predator or human observer. A recent study indicated that titmice modify the note composition of their calls in the presence of such threatening stimuli. Here, we tested whether female and male titmice differed in their calling behavior, as relatively few sex differences have been documented in calls shared by female and male songbirds. Individual titmice were captured in walk‐in treadle traps, and we gradually increased the level of fear or arousal by approaching and finally capturing the bird in the hand. Male titmice produced more chick‐a‐dee calls than females as the level of threat increased, up to the point of capturing the bird in the hand. Furthermore, the note composition of calls produced by males differed from that of calls produced by females. A limitation to our study is that our method did not allow us to rule out the possibility that size or dominance differences, rather than sex, were the main reason for the differences in calling we detected. However, increased size generally was not associated with increased calling. We discuss some possible explanations for variation in distress calling behavior in titmice.