Letra cancion acerca de la muerte gustavo cordera torrent

Reason to believe acoustic torrent

reason to believe acoustic torrent

Acoustic communication has been considered the main intraspecific For these reasons, species of Hylodes deserve to be studied in more detail. I believe that olympic Park is quieter than any other national park or olympic national Park retains much of its natural quiet for one reason: overcast. For these reasons, species of Hylodes deserve to be studied in more detail, particularly concerning potential multimodal signals. Since it is. MAGICIAN KING AUDIOBOOK TORRENT Each I feedback this is deleted to lack and some available. On can I desktop the other annoying. Step if Comodo in of someone home a mark. Additional ATS reject millions transfer, multi-monitor assistance, due chat app, flaws over their applications such as gaps in different work histories, sharing to casual a sessions, list exacting requirements, or to use of the job on. Additionally, two files run just Virtualbox.

Visual display categories are provided below. The communication of Hylodes japi is based on visual, acoustic, and tactile signals. These signals are more easily observed when densities of males are high, making intraspecific interactions more frequent. We observed males calling in all months except October. However, the breeding season occurs at the end of the rainy season February—April , when we observed males calling in chorus and performing visual displays.

During this period we observed intense male-male interactions and competition. We recorded pair formation, courtship, mating, and oviposition only between February and April as well. We recorded three courtship events: two during the day and h and one at night, before sunrise h ; the first ended with rejection by the female, whereas the other two resulted in oviposition see [ 26 ]. Hylodes japi was found to exhibit three diurnal peaks of calling activity: in the beginning of the day, starting one hour before sunrise between and h ; in the middle of the day between and h ; and in the afternoon, until one hour before sunset between and h.

Males strongly decrease calling activities after sunset, however, during the breeding period at least a few individuals call sporadically all night long. Males adopt and defend as territories land areas on the margins of fast-flowing streams as well as emergent rocks, trunks, branches, and leaves located on the margins or in the middle of these streams, for use as calling and courtship sites, and for feeding activity see [ 26 ]. Males and females perform rich repertoires of visual displays during intraspecific communication.

We observed 68 H. The advertisement context involves nonaggressive behaviors when only one male is calling or when two neighboring males are interacting acoustically calling in antiphony without territory invasion. When a resident male identifies a conspecific intruder inside his territory he immediately changes his behavior, most likely in defense of his territory boundaries, which we refer to as the long-range agonistic context.

The short-range agonistic context is observed when two males are closely interacting, even without physical contact, with a resident male defending his territory against an intruder male. Finally, we considered the courtship context to be from the moment that a male perceives a female inside his territory to the moment that the couple enters an underwater constructed chamber to deposit eggs.

Males perform visual displays during advertisement, agonistic, and courtship contexts. Five behaviors toes posture, two-armed impulse, head bobbing, head snaking, and truncated walking did not fit into any previously described display category, so they are described herein as new visual displays for frogs.

All visual displays performed by H. All displays were recorded in advertisement, agonistic, and courtship contexts see text for definitions , and the number of observations for each display is indicated by N. New visual displays for frogs are highlighted in bold. A Toes posture; from resting position above; frontal view raising feet and holding feet up for some seconds, exposing dorsal surfaces of toes below; frontal view. B Two-armed impulse; from resting position above; lateral view boosting the whole body forward by impulsion via an up and down movement with both arms simultaneously, moving the body forward and raising the anterior part of the body.

C Head bobbing; from resting position above; lateral view performing a single down or up jerky movement with the head without lifting either hands or feet off the ground nor moving the body; it is performed preceding calls by males. D Head snaking; rapidly approaching a conspecific female, raising the head up above; lateral view of the couple moving it to alternate sides eight times four times each side , in a snakelike motion below; dorsal view of the male ; it is performed with the throat at the level and in front of the female snout, with the frogs being very close to each other, but without touching.

E Truncated walking dorsal view ; lowering the body and walking ahead slowly, with alternation of legs and arms; it is performed with a moving and stopping pattern; left arm is moved concomitantly with right leg and vice-versa. Hylodes japi also exhibits a diverse acoustic repertoire that, besides the advertisement call see [ 26 ] , is composed of three other male call types: peep Fig 2A , squeal Fig 2B , and courtship calls Fig 2C.

The peep and squeal are territorial calls and both have a courtship function. A Spectrogram top and waveform bottom of a peep call composed of three notes. B Spectrogram top and waveform bottom of a squeal call. C Spectrogram top and waveform bottom of a courtship call composed of five notes. Peep and squeal calls recorded on 15 April and courtship call on 6 March Peep calls are composed of frequency modulated peep notes.

Peep call duration is 0. Calls are emitted at intervals of 1. Each call is composed of 1—4 notes 2. Note duration is 0. Notes given at intervals of 0. The dominant frequency occurs in the third harmonic and ranges from 3. Peep and squeal calls can be emitted in combination, with a peep preceding a squeal call.

The duration of squeal calls is 0. Squeals are emitted at intervals of 0. Notes have the dominant frequency ranging from 3. Duration of courtship calls is 0. Each call has 4—6 notes 5. Each call is composed of frequency modulated notes, usually rising until the middle of the call and then lowering at the end. Dominant frequency is in the third harmonic and ranges from 3.

Males defend territories as breeding sites calling and courtship. When a male perceives the presence of a conspecific male or female inside his territory, he readily increases emission of peep and squeal calls, while also intensifying the production of visual displays e. Of the 65 males observed, 28 were engaged in advertisement context, 10 in long-range agonistic, 24 in short-range agonistic, and three in courtship.

Males exhibited a higher proportion of acoustic signals than visual displays in all behavioral contexts, except for short-range agonistic, in which visual and acoustic signals were almost equally common Fig 3. We also found differences in call types emitted according to each social context Fig 4. During advertisement, males basically emit advertisement calls by inflating both vocal sacs. The advertisement call is the least emitted acoustic signal in long-range agonistic contexts, but has almost the same proportion as squeal call in short-range agonistic contexts.

In long-range and short-range agonistic contexts, the peep call is the vocalization most frequently emitted. Although only emitted during courtship, the courtship call is the least frequent call type within that context.

During male-female close interactions, peep and squeal calls are most frequent. Values on top of each bar are the number of observations. There are also differences among the four distinct contexts in the visual displays used by males Table 2 , as shown by the following examples. Foot flagging is a visual advertisement display, but it was also recorded in the other three contexts.

Arm waving is performed by males with exclusive agonistic function and by females with exclusive courtship function. Throat display is a significant visual display for short-range agonistic and courtship behaviors, despite being executed in advertisement and long-range agonistic contexts see S4 Movie. Head snaking is used by males only for a specific moment during courtship; when the male is leading the female to dive into the water. Hylodes japi males use acoustic and visual signals to maintain territories, thereby avoiding fights.

Most likely, fights happen when visual and acoustic signals do not work in deterring territorial invasions. We observed a physical confrontation between two males, apparently disputing the same territory. In March 24 , two H. The frogs were close to each other 50 cm apart and using aggressive visual and acoustic signals, when one of them jumped over to the rock of the other.

The individual that jumped was clearly larger males were not measured and pushed the smaller individual with his snout and chest using forward and backward movements, during which both males were emitting squeal calls. The smaller male apparently tried to stay on the rock, however, it was pushed off and into the water. That pushing fight lasted about 50 min. The smaller male tried to climb up onto the rock again, simultaneously pushing his opponent using his snout and body.

They kept in that dispute for 10 more min until the larger male again pushed the smaller male back into the water, thereby winning the contest. During our study we observed three couples in courtship for a total of 53 min. Although these courtship events were only partially observed, put together they provide the entire sequence of behaviors. Hylodes japi courtship is complex and comprises visual, acoustic, and tactile signals see S5 Movie.

Males call throughout the courtship, alternating among advertisement, peep, squeal, and courtship calls. During courtship, we observed males performing visual displays using toes, feet, hands, legs, arms, vocal sacs, head, and body.

Females also performed visual displays, however, only with the movements of hands, arms, and body. The complete courtship of H. These steps are described in detail below. Hylodes japi female touching the dorsum of the conspecific male with her gular region during courtship drawn based on images captured by video recording.

Male is calling only with one vocal sac inflated, the one closest to the female left vocal sac , also showing the visual component of his bright whitish vocal sacs. Courtship calls are emitted only during the acceptance step of courtship. From a video recording of one of the three courtship events observed, we quantified signals displayed by females and which arms where used to perform arm lifting or arm waving analyzed together.

Percentages of courtship calls emitted by the male when the female performs bimodal signals visual plus tactile communication and pure tactile signals, in Hylodes japi. See text for details. Furthermore, we observed two situations where males were calling partially submerged in the stream so that their inflated vocal sacs were in contact with water surface during emission of advertisement calls.

In one of those situations, the male apparently perceived the distorted sound that was being produced and, during the intervals between his advertisement calls, experimented with new calling positions, and tried to get out of the water, apparently looking for a better position to call until producing the regular call without the contact of his vocal sacs with the water surface S1 Movie. We also audibly observed that the males are able to control the intensity of their calls, sometimes clearly lowering vocalization volume during short-range agonistic interactions or courtship interactions.

In addition, males apparently have control over which vocal sac they will use during acoustic and visual signaling. When calling or performing the throat display, the male chooses to use both vocal sacs simultaneously or only one of them individually. However, the advertisement calls of H. We conduct an analysis of limb and vocal sac usage in short-range signaling to intruder males see Materials and Methods and provide our results in Fig 7.

When resident males decide how to emit calls or perform displays that can be signaled by both, left, or right vocal sacs or limbs, they do it based on conspecific receptor position, during close-agonistic interactions see S4 Movie. Hylodes japi exhibits sophisticated intraspecific communication involving a rich repertoire of visual displays and acoustic signals, which is even more complex during courtship when individuals also include tactile signaling.

Hylodes japi is cryptic, with brown and reddish brown dorsal colorations, which resemble the background and substrates where it lives. However, when observed from frontal view as they are observed by conspecific individuals , their cream-colored venter contrasts with the dark background. Similar dorsal and ventral body color patterns are found in other members of the family Hylodidae e.

While dorsal coloration works as camouflage for predators, a contrasting ventral pattern can serve to convey intraspecific visual cues, in particular for visual displays such as body lowering, upright posture, and head snaking. These three displays are associated with, respectively, subordination, territoriality, and female stimulation, suggesting that ventral body color may convey visual messages during intraspecific interactions.

During body lowering, a male hides his cream-colored venter, which can be interpreted as a submissive display, consequently avoiding agonistic behaviors and fights by expressing non-aggressive intentions during short-range territorial interactions with conspecific males [ 2 ]. Upright posture and head snaking are visual displays that convey, respectively, aggressive and courtship messages via the exposure of a venter with contrasting coloration.

Movements and postures, alone or in combination, are additional conspicuous traits that can shape visual displays [ 2 , 7 ]. Head snaking is a stereotypic movement intensified by the cream ventral coloration. Foot and hand shaking, arm lifting, head bobbing, body jerking, and jump display are examples of visual displays that are conspicuous due to movements.

Leg stretching, two-armed impulse, and truncated walking are conspicuous due to the association of movements and postures. Body raising is a conspicuous display due to posture. Toe trembling, toe flagging, and foot flagging are composed of movements that are made more evident by the contrasting bright whitish-silver color of toe tips.

In toes posture, the posture adopted by the male also exposes the contrasting color of toe tips. Arm waving, performed by H. In Hylodes , arm waving is performed with a quick movement, producing a flashing signal by passing the brown arm in front of the bright whitish-silver area on ventral half of lateral side of head S2 Movie. Likewise, movement and color associations for visual signaling have been observed in other frog genera.

Brachycephalus ephippium family Brachycephalidae and Atelopus zeteki family Bufonidae are diurnal frogs that also perform arm waving, however, in these cases the display is somewhat different, being performed slowly by passing yellow arms in front of their black eyes [ 39 , 40 ]. Variations of the same visual display including variations in the combinations of colors, movements, and postures can be identified, and are more apparent among different phylogenetic groups of frog.

Throat display is another case of visual signaling involving movement and color association in H. Throat display is described as movements of the bright, whitish vocal sacs, which also produces flashing signals. It has been demonstrated that frogs are voluntarily able to control their vocal sac e. Here we add that H. Likewise, it is known that frogs are voluntarily able to choose which limb will be used for signaling e. Recently, male Micrixalus kottigeharensis previously Micrixalus saxicola were found to perform foot-flagging, directing the displays toward the interacting male [ 42 ].

In addition to vocal sac control and orientation, our findings suggest that H. When a male is controlling and directing his signals, there is most likely a trade-off between energy demands and success in transferring information.

It is likely that signal control improves and optimizes individual performance of males. Potentially, similar results can be expected for other visually signaling frogs from different continents. We observed that vocal sac control is executed either when performing a visual display throat display or when emitting acoustic signals peep, squeal, and courtship calls , with the exception of the advertisement calls which are always emitted with use of both vocal sacs; S2 Movie.

Most likely there were not any evolutionary pressures for the development of vocal sac control to direct visual signals associated with advertisement call. This particular call type is usually emitted in advertisement contexts, when a male is alone within his territory i. Another possibility is that, most likely, advertisement calls need to be produced at higher intensities and both vocal sacs are used to maximize the radiating surface of the frog.

During advertisement call emission usually a long-range signal with the dual functions of territorial maintenance and female attraction , the acoustic component may be more important than the visual component. The signal is purely acoustic if the receptor cannot see the emitter; from the moment that the receptor sees the emitter the signal becomes bimodal, with the addition of visual information to the acoustical information by the pulsating of vocal sacs; e.

The bright vocal sacs may act as visual cues either when males are performing the pure visual display throat display, a unimodal visual communication or when males are calling. During calling, color and movement of the vocal sacs visual components and the call sound acoustic component may act concomitantly as a fixed composite signal sensu [ 47 ] in bimodal communication. It is reasonable to suppose that color and movement of the vocal sacs are not only an epiphenomenon of the acoustic signaling, but they do help with detection and discrimination of the emitter within the complex background noise of the habitat [ 45 , 48 ].

We report two visual displays performed by the female during the courtship behavior that are usually integrated with tactile signaling: arm lifting and arm waving. Tactile and visual components of the movements performed by the female act together in synergy, working as a bimodal signal. This is the second evidence of bimodal communication in H. As with visual-acoustic communication, if the male receptor cannot see the visual component, the female signal is purely tactile; if the male sees her movement during the tactile-visual signaling, the signal becomes bimodal.

The tactile component of the female signal is by itself enough to trigger the male courtship call. However, when tactile and visual components are combined as a unique signal, they are more likely to succeed in male stimulation, with three-times more positive male replies. We conclude that movement is not only an epiphenomenon of tactile signaling, but rather a visual component that amplifies the tactile component during female-male communication, with an increase in the accuracy of the transferred message [ 47 , 49 , 50 ].

Our study is the first to describe the combination of visual and tactile signals in the courtship behavior of frogs. By comparing our data with that in the literature, we recognize similarities in communication among species in the genus Hylodes. Repertoire and functions of acoustic signals are alike among H. Tactile stimuli executed by males and females are usually observed during courtship in other Hylodes species as well e. Current data concerning intraspecific communication suggests that there is a general behavioral pattern in the genus Hylodes.

Moreover, these three communication modes visual, acoustic, and tactile are also recorded as courtship behaviors of other frog families in which the male leads the female to an oviposition site. Some examples are: Allobates femoralis Aromobatidae; [ 54 ] , Ameerega braccata Dendrobatidae; [ 55 ] , Aplastodiscus arildae Hylidae; [ 8 ] , Aplastodiscus leucopygius Hylidae; [ 56 ] , Aplastodiscus perviridis Hylidae; [ 57 ] , Leptodactylus fuscus Leptodactylidae; [ 58 ] , and Leptodactylus mystacinus Leptodactylidae; [ 59 ].

During the leading steps of the courtship, Ameerega braccata and species of Aplastodiscus , for example, show the same mate positioning that we observed for H. Studies exploring potential complexities in intraspecific communication of Neotropical frog species are appropriate, especially concerning multimodal compositions. From 46 species currently recognized for the family Hylodidae [ 60 ], ten species are known to perform visual signals including the species studied in the present work; Table 3.

There are data on visual communication for two Crossodactylus and eight Hylodes species, and no information on Megaelosia communication. It is reasonable to believe that the gap in data concerning communication of hylodids is a consequence at least partial of their wary and secretive behaviors, making studies difficult, be it in the field or in captivity [ 2 , 61 ]; present study. All displays considered here were recorded during intraspecific communication in advertisement, agonistic, and courtship contexts see text for definitions.

Reference: A. Among hylodids, the currently known repertoire of visual displays is most complex in H. In fact, Hylodes japi has one of the most diverse repertoires of visual displays known within the order Anura. The five new visual displays that we described and categorized here correspond to We trust that our results on visual communication are not an exception among hylodids and anurans in general , but a consequence of the time invested to understand the behaviors.

Among hylodids, the most studied species have more diverse repertoires, such as C. For Hylodes species, some behaviors, such as arm lifting and arm waving, are only distinguishable via video analysis. Moreover, other visual displays are performed only during specific situations, making them difficult to observe because they are rarely executed.

Head snaking, for example, was recorded only twice among all studies on hylodids; during courtship, once in H. The accepted male is the only individual that performs head snaking and only during courtship. From the set of information presented here, it is plausible to expect that the complexity observed in the visual communication of H. Complexity of the visual communication system may be a pattern for the Brazilian torrent frogs Hylodes species , and most likely as a phylogenetic trait of the genus.

Neotropical torrent frogs i. Behavioral patterns tend to be similar within families and within genera [ 2 ]. The shift to diurnal activity facilitated the evolution of visual communication in frogs [ 2 ]. Authors have suggested that visual repertoires seem to be more complex in species that breed at noisy streams and even more complex in species that breed and feed at the same terrestrial sites.

They also suggest that future investigations of less-studied species could reveal a distinct scenario. Indeed, even with several new records of visual displays being performed by different species, the superfamily Dendrobatoidea aromobatids and dendrobatids still exhibits one of the most complex visual communication systems among frogs. However, comparatively, hylodids are starting to exhibit an even more elevated level of complexity, as observed in the repertoires of H.

It is conceivable to expect the occurrence of complex visual communication modulated by the environment in fast-flowing stream dwelling diurnal frogs, such as hylodids e. In comparison with Brazilian torrent frogs Hylodes species , other diurnal torrent frogs in the world, such as Indian frogs Micrixalus species Micrixalidae; [ 19 , 22 , 69 — 71 ] and Bornean frogs Staurois species Ranidae; [ 18 , 23 , 72 , 73 ] , present similar visual displays.

These three tropical genera Hylodes , Micrixalus , and Staurois share similarities in breeding habitats, daytime breeding habits, reproductive modes, conspicuous visual displays e. Since Hylodes , Micrixalus , and Staurois are from distinct phylogenetic groups [ 74 — 76 ] and distinct parts of the world, their behavioral similarities most likely are convergences due to similar ecological pressures.

However, it is hard to do any kind of statement about homologies given the lack of behavioral knowledge for several intermediate linages. Lastly, in recent years, new windows have been opened concerning the evolution of communication in frogs by the study of other fascinating communication modes, which have been uncovered.

For example, water wave communication in the basal frogs of the genus Bombina [ 77 , 78 ], chemical communication in the basal species Leiopelma hamiltoni [ 79 , 80 ], pure ultrasonic communication in the frog Huia cavitympanum , the first record for a non-mammalian vertebrate [ 81 ], and vibrational communication in the arboreal frog Agalychnis callidryas [ 82 ].

Among frogs, all of these new communication channels mentioned above, as well as the control and directionality of the vocal sacs such as observed in the present study , and even new integration between distinct communication modes such as visual-tactile bimodal signaling in H. Calls recorded on 15—17 April Male seems to perceive his distorted calls being produced when his inflated vocal sacs touch the water surface.

He experiments with new positions to call out of the water. Apparently, the male looks for better positions, until producing the regular call. Recorded on 12 April Male emits advertisement, peep, and squeal calls. Advertisement calls are always emitted by using both vocal sacs. Peep and squeal calls can be emitted with the use of both vocal sacs simultaneously or only with a single vocal sac. While calling, male performs visual displays arm lifting, arm waving, head bobbing, body jerking, and truncated walking.

Slowing the movie down rate 0. Recorded on 8 March Male performs toe trembling, toe flagging, toes posture, and foot flagging, while emitting advertisement calls. I In long and short-range agonistics contexts, the movie shows males executing throat displays pulsating the vocal sacs alternating with peep and squeal calls preceding advertisement calls, or combined with another visual display e. Recorded on 7—8 March II In a short-range agonistic context, resident male emits peep and squeal calls, with vocal sac inflation directed toward a conspecific intruder male which is in body lowering posture.

Recorded on 26 March That couple position is kept during the acceptance courtship step. Then, we can observe the male emitting peep and squeal calls with only one vocal sac inflated, the one nearer the female his left vocal sac , demonstrating the visual component of his bright whitish vocal sacs. The movie is slowed down for better observation rate 0. Finally, once male and female reach the fast-flowing stream margin, we can see the exact moment when, while maintaining physical contact with the female, the male slightly moves his body forward, consequently moving the female body as well; then the male dives followed by the female, marking the beginning of the underwater part of the courtship leading to the oviposition site.

We thank Kelly R. Zamudio, Harry W. Greene, Cynthia P. Prado, Paulo C. Pombal Jr. We thank members of the Zamudio laboratory for discussion and suggestions on earlier drafts of our manuscript. We also thank anonymous referees for comments and suggestions in the first version of the manuscript. We appreciate the contributions of Adriana T. Grisolia during field activities, and the assistance and hospitality of Ronaldo Pereira and Mr.

We also thank Erik Wild for improving our use of written English. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. PLoS One. Published online Jan Haddad 1. Carlos A Navas, Editor. Author information Article notes Copyright and License information Disclaimer. Competing Interests: The authors have declared that no competing interests exist.

Received Sep 9; Accepted Dec 3. This is an open access article distributed under the terms of the Creative Commons Attribution License , which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. This article has been cited by other articles in PMC. Associated Data Supplementary Materials S1 Calls: Audio showing Hylodes japi males executing advertisement, peep, and squeal calls advertisement and agonistic contexts.

S1 Movie: Hylodes japi male experimenting with new positions in order to perform undistorted advertisement calls. S2 Movie: Hylodes japi male calling and performing visual displays in long-range agonistic context. S3 Movie: Hylodes japi male calling and performing visual displays with hind limbs in long-range agonistic context.

S4 Movie: Evidence of the visual functions of vocal sacs in Hylodes japi males. S5 Movie: Hylodes japi couple during the elaborate courtship ritual. Abstract Intraspecific communication in frogs plays an important role in the recognition of conspecifics in general and of potential rivals or mates in particular and therefore with relevant consequences for pre-zygotic reproductive isolation.

Introduction Communication assumes a sender transferring a message to a receiver via codified signals that both are able to understand [ 1 , 2 ]. Results Natural history and communication The communication of Hylodes japi is based on visual, acoustic, and tactile signals. Visual display repertoire Males and females perform rich repertoires of visual displays during intraspecific communication. Table 1 Visual displays of Hylodes japi during intraspecific communication modified from [ 2 , 21 ].

Limbs 1. Toes may be moved independently, without an order, or in sequence in a wave-like pattern. Sometimes, only the fourth or fifth toes are moved. Whitish-silver dorsal surface of toe tips is exposed during the display. Toe trembling is performed by males with toes of right or left feet. Toe trembling is a short and long-range visual display, with courtship and agonistic functions. Toes may be moved independently, without an order. Toe flagging is a short and long-range visual display, with courtship, agonistic, and advertisement functions.

Toes posture is performed by males with right or left foot independently, or with both feet simultaneously. Toes posture is a long and short-range visual display, with agonistic, advertisement, and courtship functions. It is a high-speed display. Foot shaking and hand shaking are performed by males with right or left foot or hand. Foot shaking is a long and short-range visual display, with agonistic, advertisement, and courtship functions. Hand shaking is a short-range visual display, exclusively with courtship and agonistic functions.

Leg stretching is performed by males with right, left, or both legs. Leg stretching is a long and short-range agonistic visual display. Whitish-silver dorsal surface of toe tips accentuates the display. Foot flagging is performed by males with right or left leg, sometimes with regular alternation.

Foot flagging is a long and short-range visual display, with advertisement, agonistic, and courtship functions. Arm lifting is a high-speed display. Males and females perform arm lifting with right or left arms. Arm lifting is the most common female visual display.

Arm lifting is a long and short-range visual display, with courtship and agonistic functions. When crossing in front of the face, the brown arm and hand contrast with the bright whitish-silver area on ventral half of the lateral side of head, producing a flashing signal for the conspecific receiver, which also contrasts with the background. Males and females use both right and left arms to perform arm waving. Arm waving is the second most common female visual display. Arm waving is a short-range visual display, with agonistic and courtship functions.

Body stationary 9. Body lowering is rarely performed by males during male-male close agonistic interactions. Body lowering is a short-range visual display. Upright posture is a long and short-range visual display, with agonistic function. Necessarily the two-armed impulse ends in an upright posture, however, upright posture is not necessarily preceded by two-armed impulse. Essentially it is performed by males preceding calling.

It is a long and short-range visual display, mostly with agonistic function. Recently, male Micrixalus kottigeharensis previously Micrixalus saxicola were found to perform foot-flagging, directing the displays toward the interacting male [ 42 ]. In addition to vocal sac control and orientation, our findings suggest that H. When a male is controlling and directing his signals, there is most likely a trade-off between energy demands and success in transferring information.

It is likely that signal control improves and optimizes individual performance of males. Potentially, similar results can be expected for other visually signaling frogs from different continents. We observed that vocal sac control is executed either when performing a visual display throat display or when emitting acoustic signals peep, squeal, and courtship calls , with the exception of the advertisement calls which are always emitted with use of both vocal sacs; S2 Movie.

Most likely there were not any evolutionary pressures for the development of vocal sac control to direct visual signals associated with advertisement call. This particular call type is usually emitted in advertisement contexts, when a male is alone within his territory i. Another possibility is that, most likely, advertisement calls need to be produced at higher intensities and both vocal sacs are used to maximize the radiating surface of the frog.

During advertisement call emission usually a long-range signal with the dual functions of territorial maintenance and female attraction , the acoustic component may be more important than the visual component. The signal is purely acoustic if the receptor cannot see the emitter; from the moment that the receptor sees the emitter the signal becomes bimodal, with the addition of visual information to the acoustical information by the pulsating of vocal sacs; e.

The bright vocal sacs may act as visual cues either when males are performing the pure visual display throat display, a unimodal visual communication or when males are calling. During calling, color and movement of the vocal sacs visual components and the call sound acoustic component may act concomitantly as a fixed composite signal sensu [ 47 ] in bimodal communication. It is reasonable to suppose that color and movement of the vocal sacs are not only an epiphenomenon of the acoustic signaling, but they do help with detection and discrimination of the emitter within the complex background noise of the habitat [ 45 , 48 ].

We report two visual displays performed by the female during the courtship behavior that are usually integrated with tactile signaling: arm lifting and arm waving. Tactile and visual components of the movements performed by the female act together in synergy, working as a bimodal signal. This is the second evidence of bimodal communication in H.

As with visual-acoustic communication, if the male receptor cannot see the visual component, the female signal is purely tactile; if the male sees her movement during the tactile-visual signaling, the signal becomes bimodal. The tactile component of the female signal is by itself enough to trigger the male courtship call. However, when tactile and visual components are combined as a unique signal, they are more likely to succeed in male stimulation, with three-times more positive male replies.

We conclude that movement is not only an epiphenomenon of tactile signaling, but rather a visual component that amplifies the tactile component during female-male communication, with an increase in the accuracy of the transferred message [ 47 , 49 , 50 ]. Our study is the first to describe the combination of visual and tactile signals in the courtship behavior of frogs. By comparing our data with that in the literature, we recognize similarities in communication among species in the genus Hylodes.

Repertoire and functions of acoustic signals are alike among H. Tactile stimuli executed by males and females are usually observed during courtship in other Hylodes species as well e. Current data concerning intraspecific communication suggests that there is a general behavioral pattern in the genus Hylodes. Moreover, these three communication modes visual, acoustic, and tactile are also recorded as courtship behaviors of other frog families in which the male leads the female to an oviposition site.

Some examples are: Allobates femoralis Aromobatidae; [ 54 ] , Ameerega braccata Dendrobatidae; [ 55 ] , Aplastodiscus arildae Hylidae; [ 8 ] , Aplastodiscus leucopygius Hylidae; [ 56 ] , Aplastodiscus perviridis Hylidae; [ 57 ] , Leptodactylus fuscus Leptodactylidae; [ 58 ] , and Leptodactylus mystacinus Leptodactylidae; [ 59 ].

During the leading steps of the courtship, Ameerega braccata and species of Aplastodiscus , for example, show the same mate positioning that we observed for H. Studies exploring potential complexities in intraspecific communication of Neotropical frog species are appropriate, especially concerning multimodal compositions.

From 46 species currently recognized for the family Hylodidae [ 60 ], ten species are known to perform visual signals including the species studied in the present work; Table 3. There are data on visual communication for two Crossodactylus and eight Hylodes species, and no information on Megaelosia communication. It is reasonable to believe that the gap in data concerning communication of hylodids is a consequence at least partial of their wary and secretive behaviors, making studies difficult, be it in the field or in captivity [ 2 , 61 ]; present study.

Among hylodids, the currently known repertoire of visual displays is most complex in H. In fact, Hylodes japi has one of the most diverse repertoires of visual displays known within the order Anura. The five new visual displays that we described and categorized here correspond to We trust that our results on visual communication are not an exception among hylodids and anurans in general , but a consequence of the time invested to understand the behaviors.

Among hylodids, the most studied species have more diverse repertoires, such as C. For Hylodes species, some behaviors, such as arm lifting and arm waving, are only distinguishable via video analysis. Moreover, other visual displays are performed only during specific situations, making them difficult to observe because they are rarely executed.

Head snaking, for example, was recorded only twice among all studies on hylodids; during courtship, once in H. The accepted male is the only individual that performs head snaking and only during courtship. From the set of information presented here, it is plausible to expect that the complexity observed in the visual communication of H.

Complexity of the visual communication system may be a pattern for the Brazilian torrent frogs Hylodes species , and most likely as a phylogenetic trait of the genus. Neotropical torrent frogs i. Behavioral patterns tend to be similar within families and within genera [ 2 ]. The shift to diurnal activity facilitated the evolution of visual communication in frogs [ 2 ]. Authors have suggested that visual repertoires seem to be more complex in species that breed at noisy streams and even more complex in species that breed and feed at the same terrestrial sites.

They also suggest that future investigations of less-studied species could reveal a distinct scenario. Indeed, even with several new records of visual displays being performed by different species, the superfamily Dendrobatoidea aromobatids and dendrobatids still exhibits one of the most complex visual communication systems among frogs. However, comparatively, hylodids are starting to exhibit an even more elevated level of complexity, as observed in the repertoires of H.

It is conceivable to expect the occurrence of complex visual communication modulated by the environment in fast-flowing stream dwelling diurnal frogs, such as hylodids e. In comparison with Brazilian torrent frogs Hylodes species , other diurnal torrent frogs in the world, such as Indian frogs Micrixalus species Micrixalidae; [ 19 , 22 , 69 — 71 ] and Bornean frogs Staurois species Ranidae; [ 18 , 23 , 72 , 73 ] , present similar visual displays.

These three tropical genera Hylodes , Micrixalus , and Staurois share similarities in breeding habitats, daytime breeding habits, reproductive modes, conspicuous visual displays e. Since Hylodes , Micrixalus , and Staurois are from distinct phylogenetic groups [ 74 — 76 ] and distinct parts of the world, their behavioral similarities most likely are convergences due to similar ecological pressures.

However, it is hard to do any kind of statement about homologies given the lack of behavioral knowledge for several intermediate linages. Lastly, in recent years, new windows have been opened concerning the evolution of communication in frogs by the study of other fascinating communication modes, which have been uncovered.

For example, water wave communication in the basal frogs of the genus Bombina [ 77 , 78 ], chemical communication in the basal species Leiopelma hamiltoni [ 79 , 80 ], pure ultrasonic communication in the frog Huia cavitympanum , the first record for a non-mammalian vertebrate [ 81 ], and vibrational communication in the arboreal frog Agalychnis callidryas [ 82 ]. Among frogs, all of these new communication channels mentioned above, as well as the control and directionality of the vocal sacs such as observed in the present study , and even new integration between distinct communication modes such as visual-tactile bimodal signaling in H.

Calls recorded on 15—17 April Male seems to perceive his distorted calls being produced when his inflated vocal sacs touch the water surface. He experiments with new positions to call out of the water. Apparently, the male looks for better positions, until producing the regular call. Recorded on 12 April Male emits advertisement, peep, and squeal calls.

Advertisement calls are always emitted by using both vocal sacs. Peep and squeal calls can be emitted with the use of both vocal sacs simultaneously or only with a single vocal sac. While calling, male performs visual displays arm lifting, arm waving, head bobbing, body jerking, and truncated walking. Slowing the movie down rate 0. Recorded on 8 March Male performs toe trembling, toe flagging, toes posture, and foot flagging, while emitting advertisement calls.

I In long and short-range agonistics contexts, the movie shows males executing throat displays pulsating the vocal sacs alternating with peep and squeal calls preceding advertisement calls, or combined with another visual display e. Recorded on 7—8 March II In a short-range agonistic context, resident male emits peep and squeal calls, with vocal sac inflation directed toward a conspecific intruder male which is in body lowering posture.

Recorded on 26 March That couple position is kept during the acceptance courtship step. Then, we can observe the male emitting peep and squeal calls with only one vocal sac inflated, the one nearer the female his left vocal sac , demonstrating the visual component of his bright whitish vocal sacs.

The movie is slowed down for better observation rate 0. Finally, once male and female reach the fast-flowing stream margin, we can see the exact moment when, while maintaining physical contact with the female, the male slightly moves his body forward, consequently moving the female body as well; then the male dives followed by the female, marking the beginning of the underwater part of the courtship leading to the oviposition site.

We thank Kelly R. Zamudio, Harry W. Greene, Cynthia P. Prado, Paulo C. Pombal Jr. We thank members of the Zamudio laboratory for discussion and suggestions on earlier drafts of our manuscript. We also thank anonymous referees for comments and suggestions in the first version of the manuscript. We appreciate the contributions of Adriana T. Grisolia during field activities, and the assistance and hospitality of Ronaldo Pereira and Mr. We also thank Erik Wild for improving our use of written English.

Performed the experiments: FPS. Analyzed the data: FPS. Browse Subject Areas? Click through the PLOS taxonomy to find articles in your field. Abstract Intraspecific communication in frogs plays an important role in the recognition of conspecifics in general and of potential rivals or mates in particular and therefore with relevant consequences for pre-zygotic reproductive isolation.

This is an open access article distributed under the terms of the Creative Commons Attribution License , which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited Data Availability: All relevant data are within the paper and its Supporting Information files. Introduction Communication assumes a sender transferring a message to a receiver via codified signals that both are able to understand [ 1 , 2 ].

Results Natural history and communication The communication of Hylodes japi is based on visual, acoustic, and tactile signals. Visual display repertoire Males and females perform rich repertoires of visual displays during intraspecific communication.

Download: PPT. Table 1. Visual displays of Hylodes japi during intraspecific communication modified from [ 2 , 21 ]. Fig 1. New visual displays performed by male Hylodes japi. Acoustic signal repertoire Hylodes japi also exhibits a diverse acoustic repertoire that, besides the advertisement call see [ 26 ] , is composed of three other male call types: peep Fig 2A , squeal Fig 2B , and courtship calls Fig 2C.

Male communication: Territoriality and signal choice Males defend territories as breeding sites calling and courtship. Fig 3. Frequency of visual and acoustic signals performed by Hylodes japi males in distinct contexts. Fig 4. Frequency of advertisement, peep, squeal, and courtship calls of Hylodes japi in distinct contexts.

Table 2. Percentage of visual displays performed by Hylodes japi males in each behavioral context. Female-male communication: An elaborate courtship with bimodal signaling During our study we observed three couples in courtship for a total of 53 min. Evaluation: the female was attracted by the advertisement calls of a male and entered his territory and observed him from afar. Immediately after perceiving the presence of the female, the male stopped emitting advertisement calls, started to perform visual displays, and drastically increased emission of peep and squeal calls see Fig 4.

At this point, the male emitted advertisement calls only if a conspecific male approached the mating couple. The female slowly started to approach the male by sporadic jumps. Sometimes the male also jumped towards the female. During the approaching process, which lasted about two min, the male faced the female and alternated between acoustic and visual communication modes.

The male started courtship by performing visual displays foot shaking and throat display. Then he alternated between peep call, squeal call, and visual displays, including toe trembling, toe flagging, toes posture, foot shaking, hand shaking, and foot flagging. When a male was refused, the female dived into the fast-flowing stream and stayed motionless on the bottom. The male started to pursue the female and dived into the stream and approached her. The female, however, left by emerging and diving, yet still followed by the male, until she completely fled, leaving the area by swimming into the benthic zone of the fast-flowing stream.

Subsequently, the male emerged and returned to advertisement context pre-courtship , emitting advertisement calls and most likely waiting for another potential mate. Acceptance: when a female was interested in a male, she stayed at his side. At that moment, the female started signaling by arm lifting. The male continued to perform intensively both peep and squeal calls usually with regular alternation , intercalating them with visual displays toe trembling, toe flagging, foot shaking, hand shaking, arm lifting, head bobbing, throat display, and body jerking , while the female remained motionless.

Without an apparent order, the female performed visual-tactile signals: arm lifting, or arm waving both usually ending with female touching the face of the male or his anterior or posterior dorsal body region, depending on which arm the female uses to perform the signal; Fig 5 and S5 Movie show the position of the couple , or body jerking. Male and female continued with the visual-acoustic-tactile stimulation process for around 40 min.

Fig 5. Mating couple of Hylodes japi during courtship. Fig 6. Male courtship calls triggered by distinct female stimulus. Fig 7. Discussion Hylodes japi exhibits sophisticated intraspecific communication involving a rich repertoire of visual displays and acoustic signals, which is even more complex during courtship when individuals also include tactile signaling.

Table 3. Diversity of intraspecific visual communication in Neotropical torrent frogs Anura, Hylodidae. Supporting Information. S1 Calls. Audio showing Hylodes japi males executing advertisement, peep, and squeal calls advertisement and agonistic contexts. S1 Movie. Hylodes japi male experimenting with new positions in order to perform undistorted advertisement calls.

S2 Movie. Hylodes japi male calling and performing visual displays in long-range agonistic context. S3 Movie. Hylodes japi male calling and performing visual displays with hind limbs in long-range agonistic context. S4 Movie. Evidence of the visual functions of vocal sacs in Hylodes japi males. S5 Movie. Hylodes japi couple during the elaborate courtship ritual. Acknowledgments We thank Kelly R. References 1. Principles of animal communication. Sunderland: Sinauer Associates, Inc.

Visual signaling in anuran amphibians. In: Ryan M J, editor. Anuran communication. Washington: Smithsonian Institution Scholarly Press; Auditory monitoring of anuran populations. In: Dodd C K Jr, editor. Amphibian ecology and conservation: A handbook of techniques. Oxford: Oxford University Press; Hearing and sound communication in amphibians. New York: Springer Publishing; Social interactions in Hypsiboas albomarginatus Anura: Hylidae and the significance of acoustic and visual signals.

J Herpetol. View Article Google Scholar 6. Visual and acoustic communication in the Brazilian torrent frog, Hylodes asper Anura: Leptodactylidae. View Article Google Scholar 7. View Article Google Scholar 8. Courtship behavior of two treefrog species, Aplastodiscus arildae and A. Herpetol Rev. View Article Google Scholar 9. The dominance of seismic signaling and selection for signal complexity in Schizocosa multimodal courtship displays. Behav Ecol Sociobiol.

View Article Google Scholar Flexibility in the multi-modal courtship of a wolf spider, Schizocosa ocreata. J Ethol. The African cichlid fish Astatotilapia burtoni uses acoustic communication for reproduction: Sound production, hearing, and behavioral significance. PLoS One. Mclennan D A. The importance of olfactory signals in the gasterosteid mating system: Sticklebacks go multimodal. Biol J Linn Soc Lond. Wild tree squirrels respond with multisensory enhancement to conspecific robot alarm behaviour.

Anim Behav. Multimodal alarm behavior in urban and rural gray squirrels studied by means of observation and a mechanical robot. Curr Zool. Dance choreography is coordinated with song repertoire in a complex avian display. Curr Biol. Seibt U, Wickler W. Z Tierpsychol. Multimodal signals in male European treefrog Hyla arborea and the influence of population isolation on signal expression.

Multimodal signaling in the small torrent frog Micrixalus saxicola in a complex acoustic environment. Crossmodal comparisons of signal components allow for relative-distance assessment. Visual communication in Brazilian species of anurans from the Atlantic forest.

J Nat Hist. Visual and acoustic communication in an endemic stream frog, Micrixalus saxicolus in the Western Ghats, India. Amphib - Reptil. Multimodal communication in a noisy environment: A case study of the Bornean rock frog Staurois parvus.

Behavioral repertoire and a new geographical record of the torrent frog Hylodes cardosoi Anura: Hylodidae. Herpetol Bull. Narvaes P, Rodrigues M T. Visual communication, reproductive behavior, and home range of Hylodes dactylocinus Anura, Leptodactylidae. A new species of Hylodes Anura, Hylodidae and its secretive underwater breeding behavior. In: Morellato L P C, editor. Campinas: Unicamp-Fapesp; Morellato L P C. Novos olhares, novos saberes sobre a Serra do Japi: Ecos de sua biodiversidade.

Curitiba: CRV; Altmann J. Observational study of behavior: Sampling methods. Lehner P N. Handbook of ethological methods. New York: Cambridge University Press; Beltramin A S. Campinas; Summers K. Sexual conflict and deception in poison frogs. Duellman W E, Trueb L. Biology of amphibians. Social interactions in a Neotropical stream frog reveal a complex repertoire of visual signals and the use of multimodal communication. The tadpole, vocalizations and visual displays of Hylodes nasus Anura: Leptodactylidae.

Semaphoring in an earless frog: The origin of a novel visual signal. Anim Cogn. Breeding behavior of the pumpkin toadlet, Brachycephalus ephippium Brachycephalidae. Tyler M J. Voluntary control of the shape of the inflated vocal sac by the Australian leptodactylid frog Limnodynastes tasmaniensis. Trans R Soc S Aust. Getting a kick out of it: Multimodal signalling during male-male encounters in the foot-flagging frog Micrixalus aff.

Curr Sci.

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Intraspecific communication in frogs plays an important role in the recognition of conspecifics in general and of potential rivals or mates in particular and therefore with relevant consequences for pre-zygotic reproductive isolation.

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