Risks of multimodal signaling

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Bat predators attend to dynamic
motion in frog sexual displays
Wouter Halfwerk, Marjorie M. Dixon, Kristina J. Ottens , Ryan
C. Taylor, Michael J. Ryan, Rachel A. Page and Patricia L.
Jones

OBJECTIVES:
• To determine if the fringe-lipped bat (Trichops cirrhosus) detect and
select prey (Tungara frog, Physalaemus pustulosus) by its vocal sac,
• To determine if T.cirrhosus prefer dynamic vocal sacs over static ones,
or vise-versa,
• To examine the sensory system (both echolocation and vision) used by
T.cirrhosus to detect the vocal sac in a cue isolation experiment; and
• To quantify the detection limits of T.cirrhosus visual and echolocation
system.

• Complex dance rituals • Nuptial gifts • Plumage displays • Songs and Calls • Used by females to assess the desirability of the male as a contributor to the genes of the progeny .INTRODUCTION: • Courtship requires animals to produce complex signals which favors reproductive success.

• Variation in call complexity determines reproductive success.INTRODUCTION: • Tungara Frog (Physalaemus pustulosus) • Commonly distributed on the Mexican tropics and in Northern South America • Males produce advertisement calls to attract females. . Calls are usually accompanied with vocal sac movement to achieve a novel response.

right) whine plus three chucks. The figure illustrates calls of varying complexity of the same produce whines with more chucks. INTRODUCTION: • Tungara frogs produce calls with two components. a frequency- modulated whine which can be produced by itself (simple call) or followed by 1-7 harmonic burst or chucks (complex calls). left) whine plus two chucks. right) whine plus chuck. • One chuck apparently increases the attractiveness of a male fivefold Top. and (bottom. left) whine. blue illustrations are waveforms and bottom gray- thus male frogs preference to scale illustrations are spectrograms. . (top. male: (top. (bottom.

which “eavesdrops” to its courtship rituals with the female.INTRODUCTION: • However. • Thus. males are not always observed to produce whines with chuck. • This complex behavioral response is influenced by natural and sexual selection pressures acting on the individual. males would do a simple call and a complex call alternatively. . the Fringe-lipped bat. Why? • Acoustic as well as visual signals released by male Tungara frogs is not only received by its intended receiver (female Tungara) but also by its predator. changing them based on the present risk of predation.

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and along with the dynamics of the vocal sac. • Thus.INTRODUCTION: • Moreover. increasing the attractiveness of the male to the female. . it is also of interest if multimodal signals are used by the predator (Trichops cirrhosus) to assess the prey quality. Such incorporation of a signal component into a multimodal sexual display increase the accuracy of signal transfer or enhance overall signal efficacy. creates a multimodal cue. production of the sound incidentally creates ripples on the water surface.

METHODOLOGY: Experiment 1: • Test whether bat perceive vocal sac and if dynamic vocal sac was preferred over static ones • Two model frogs was used. one which emits both acoustic cues and vocal cues while the other one emits acoustic cues only. . cirrhosus. • Both model were situated 5 meters from the perch of T.

Both the experimental catheter mimicking the túngara frog vocal sac as well as the catheter controlling for sound produced during catheter inflation are shown in the enlargement.Experimental set-up of the two choice tests. Side view of the experimental setup showing the speaker used to broadcast acoustic call component with a Plexiglas screen plus frog model on top. (A). (B) Front view showing the robofrog with inflated vocal sac on the right and the control model on the left. .

3–5  away 150 ms after peak amplitude of the from the experimental platform. Top view showing the two frog models. On the 200 ms after sound and roughly right side is the bat on the perch. Stimulus presentation showing sound playback (whine plus chuck) in the top channel and the inflation–deflation of the vocal sac in the bottom channel. whine. the Plexiglas screen and the holes underneath the Maximum inflation was reached models to allow for sound transfer. .Experimental set-up of the two choice tests.

sound off on leaving perch.METHODOLOGY: Experiment 2: • Variation of stimulus presentation • Presentation of vocal sac. vocal sac off on leaving perch • Four treatments: • Continuous • Perch only • Dynamic • Static .

METHODOLOGY: Experiment 3: • Use of echolocation to detect vocal sac was examined • Four treatments: • Visual cue only • Echolocation cue only • Both visual and echolocation cue • Visual and echolocation cue absent .

N=10. z-score=2. .RESULTS: Experiment 1: Naive Responses Wild-caught bats were given a choice to attack a frog model with a dynamic vocal sac present (inflating–deflating in synchrony with sound) or a control model (with a deflated vocal sac).79. intercept=1. All bats made their very first attack on the model with the dynamic moving vocal sac [generalized linear mixed model (GLMM).005)]. P=0.

the average attack rate on the vocal sac model was 56% higher than that of the control model. which was not attacked by the bat. Furthermore.RESULTS: Experiment 2: Effect of Stimulus Presentation The presentation of the sound and the vocal sac was varied to test whether bats perceived the vocal sac from their perch and whether the vocal sac had to be dynamically inflated and deflated. the attack rate depended on the type of presentation— there is a high preference on the dynamic inflating-deflating sac compared to static inflated sac. Sound and vocal sac were continuously presented until a bat flew from its perch towards the frog models. . Across all treatments.

RESULTS: .

n. **P<0. *P<0.001. Bats preferred to hover over and attack the frog model with a dynamically inflated–deflated vocal sac under all playback conditions. not significant.RESULTS: Bats prefer to attack a frog model with a dynamically moving vocal sac. P<0.05. . Dashed line indicates chance level at 50%..1. Attack preference did not differ from chance with a continuous inflated vocal sac (static treatment).s. ***P<0.01. Shown are boxplots derived from the model estimates per playback treatment (Experiment 2).

bats showed a significant attack preference for the vocal sac when both echolocation and visual cues were present as well as when trials were carried out in complete darkness so only echolocation cues were present. . bats showed no preference when echolocation was blocked by visually transparent fields or when both visual and echolocation cues were absent. During this experiment. However.RESULTS: Experiment 3: Use of Echolocation to Detect Vocal Sac The bats were presented with dynamic vocal sac playback and the sensory environment was manipulated to test what sensory system was used.

n.RESULTS: The cue isolation experiment reveals that bats use echolocation to detect vocal sac movements. The control condition refers to the treatment in which bats had no access to visual or echolocation cues. but not when they only had access to visual cues.s. . Shown are boxplots derived from the model estimates per cue isolation treatment (Experiment 3). Filled circles are outliers.. Bats preferred to attack the robofrog when they had only access to echolocation cues coming from the vocal sac. not significant.

Attack preference depends on the dynamic movement of the vocal sac and not just size alone. • The cue isolation experiment showed that bats used echolocation and not visual cues to detect the frog’s vocal sac.CONCLUSION: • Bats have very high preference on frogs with a vocal sac that was dynamically inflated and deflated in synchrony with acoustic call production. Thus. because bats will not attack static inflated frog models nor the static deflated ones. . the predation pressure imposed on the frog’s multimodal signal will depend on the sensory conditions that will affect bat’s echolocation.

since female tungara frogs assess the male’s vocal sac in the visual domain. . whereas the bat monitor frog vocals using echolocation.CONCLUSION: • Sexual and natural selection pressures on the same trait are not always mediated through the same sensory modalities.

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