Multimodal signaling in the Small Torrent Frog (Micrixalus saxicola) in a complex acoustic environment.

Many animals use multimodal (both visual and acoustic) components in courtship signals. The acoustic communication of anuran amphibians can be masked by the presence of environmental background noise, and multimodal displays may enhance receiver detection in complex acoustic environments. In the present study, we measured sound pressure levels of concurrently calling males of the Small Torrent Frog (Micrixalus saxicola) and used acoustic playbacks and an inflatable balloon mimicking a vocal sac to investigate male responses to controlled unimodal (acoustic) and multimodal (acoustic and visual) dynamic stimuli in the frogs' natural habitat. Our results suggest that abiotic noise of the stream does not constrain signal detection, but males are faced with acoustic interference and masking from conspecific chorus noise. Multimodal stimuli elicited greater response from males and triggered significantly more visual signal responses than unimodal stimuli. We suggest that the vocal sac acts as a visual cue and improves detection and discrimination of acoustic signals by making them more salient to receivers amidst complex biotic background noise

Testosterone Amplifies the Negative Valence of an Agonistic Gestural Display by Exploiting Receiver Perceptual Bias

Many animals communicate by performing elaborate displays that are incredibly extravagant and wildly bizarre. So, how do these displays evolve? One idea is that innate sensory biases arbitrarily favour the emergence of certain display traits over others, leading to the design of an unusual display. Here, we study how physiological factors associated with signal production influence this process, a topic that has received almost no attention. We focus on a tropical frog, whose males compete for access to females by performing an elaborate waving display. Our results show that sex hormones like testosterone regulate specific display gestures that exploit a highly conserved perceptual system, evolved originally to detect \u27dangerous\u27 stimuli in the environment. Accordingly, testosterone makes certain gestures likely to appear more perilous to rivals during combat. This suggests that hormone action can interact with effects of sensory bias to create an evolutionary optimum that guides how display exaggeration unfolds

Waste Disposal Sites as All-You-Can Eat Buffets for Carrion Crows (Corvus corone).

In cities and densely populated areas, several corvid species are considered nuisance animals. In Austria, particularly carrion (Corvus corone) and hooded crows (C. cornix) are regarded as pests by the general public that frequently cause damage to crops, feed on human waste, and thus spread trash. We conducted a detailed one-year field survey to estimate the abundance of carrion crows in relation to potential anthropogenic food sources and reference sites in the Austrian Rhine valley. Our results demonstrated that the number and proximity of waste management facilities, animal feeding areas, and agricultural areas, and the productive capacity of agricultural areas, predominantly influenced habitat choice and abundance of carrion crows. In the current study, the probability of observing more than two carrion crows at a survey site decreased with increasing human population density. Moreover, the abundance of crows increased despite a continuous increase in crow hunting kills registered during the past 25 years. Our study suggests a regionally comprehensive waste management plan could serve as a promising strategy to manage nuisance birds. A reduction in anthropogenic food supply through improved waste management practices is required for long-term, sustainable management to limit the abundance of crow populations in and close to urban environments.This research was funded by Inatura—Naturschau Dornbirn GmbH with the follwing project “name “Populationsrelevante Einflüsse von Biogas-, Grünmüllanlagen und ehemaligen Deponien auf die Corvidenfauna im Vorarlberger Rheintal: historische und rezente Entwicklungen” and was granted to Markus Boeckle

Increased Androgenic Sensitivity in the Hind Limb Muscular System Marks the Evolution of a Derived Gestural Display

Physical gestures are prominent features of many species’ multi- modal displays, yet how evolution incorporates body and leg movements into animal signaling repertoires is unclear. Andro- genic hormones modulate the production of reproductive signals and sexual motor skills in many vertebrates; therefore, one possi- bility is that selection for physical signals drives the evolution of androgenic sensitivity in select neuromotor pathways. We exam- ined this issue in the Bornean rock frog (Staurois parvus, family: Ranidae). Males court females and compete with rivals by per- forming both vocalizations and hind limb gestural signals, called “foot flags.” Foot flagging is a derived display that emerged in the ranids after vocal signaling. Here, we show that administration of testosterone (T) increases foot flagging behavior under seminatural conditions. Moreover, using quantitative PCR, we also find that adult male S. parvus maintain a unique androgenic phenotype, in which androgen receptor (AR) in the hind limb musculature is expressed at levels ∼10× greater than in two other anuran species, which do not produce foot flags (Rana pipiens and Xenopus laevis). Finally, because males of all three of these species solicit mates with calls, we accordingly detect no differences in AR expression in the vocal apparatus (larynx) among taxa. The results show that foot flagging is an androgen-dependent gestural signal, and its emergence is associated with increased androgenic sensitivity within the hind limb musculature. Selection for this novel gestural signal may therefore drive the evolution of increased AR expres- sion in key muscles that control signal production to support adap- tive motor performance

Sexual dimorphism in postcloacal scales in the northern caiman lizard (Dracaena guianensis)

Morphological differences between males and females are common among reptiles. A particularly interesting sexually dimorphic feature whose function is largely unknown is the number and pattern of specific scales. Several lizard species possess an arrangement of centered scales near the cloacal region that differ between the sexes and can be used for sex determination. The presence of postcloacal buttons, sexually dimorphic postcloacal scales on both sides of the body, is an exclusive trait in the subfamily Tupinambinae and is only poorly documented. Here, we investigate postcloacal scales in northern caiman lizards (Dracaena guianensis) housed at the Vienna Zoo. For a period of two years, we documented scale patterns and performed morphometric measurements of individuals of different age classes. Caiman lizards were CT scanned to confirm the sexes. Males exhibit three raised postcloacal scales in a row behind the left and right leg, while females possess one or two large scales surrounded by several smaller scales. The study provides the first evidence that these scales can function as a reliable trait to distinguish the sexes regardless of age or reproductive status. The sexually dimorphic bilateral scale pattern is present immediately after hatching and does not change during development. Scales only increase in thickness and length during growth. We further demonstrate that sexual size dimorphism (SSD) exists in juveniles during ontogenetic development. Juvenile females had a larger SVL, body length, tail length and higher weight compared to juvenile males. This SSD could not be confirmed in adults, and sex determination based on SSD seems unreliable

Insight into the Evolution of Anuran Foot Flag Displays: A Comparative Study of Color and Kinematics

Understanding how complex animal displays evolve is a major goal of evolutionary organismal biology. Here, we study this topic by comparing convergently evolved gestural displays in two unrelated species of frog (Bornean Rock Frog, Staurois parvus, and Kottigehara Dancing Frog, Micrixalus kottigeharensis). This behavior, known as a foot flag, is produced when a male ?waves\u27 his hindlimb at another male during bouts of competition for access to mates. We assess patterns of variation in the color of frog feet and the kinematics of the display itself to help pinpoint similarities and differences of the visual signal elements. We find clear species differences in the color of foot webbing, which is broadcast to receivers during specific phases of the display. Analyses of foot-trajectory duration and geometry also reveal clear species differences in display speed and shape - S. parvus generates a faster and more circular visual signal, while M. kottigeharensis generates a much slower and more elliptical one. These data are consistent with the notion that color, speed, and shape likely encode species identity. However, we also found that foot flag speed shows significant among-individual variation, particularly the phase of the display in which foot webbings are visible. This result is consistent with the idea that frogs alter temporal signal components, which may showcase individual condition, quality, or motivation. Overall, our comparative study helps elucidate the variability of foot flagging behavior in a manner that informs how we understand the design principles that underlie its function as a signal in intraspecific communication

Androgen Receptor Modulates Multimodal Displays in the Bornean Rock Frog (Staurois parvus)

Multimodal communication is common in the animal kingdom. It occurs when animals display by stimulating two or more receiver sensory systems, and often arises when selection favors multiple ways to send messages to conspecifics. Mechanisms of multimodal display behavior are poorly understood, particularly with respect to how animals coordinate the production of different signals. One important question is whether all components in a multimodal display share an underlying physiological basis, or whether different components are regulated independently. We investigated the influence of androgen receptors (ARs) on the production of both visual and vocal signal components in the multimodal display repertoire of the Bornean rock frog (Staurois parvus). To assess the role of AR in signal production, we treated reproductively active adult males with the antiandrogen flutamide (FLUT) and measured the performance of each component signal in the multimodal display. Our results show that blocking AR inhibited the production of multiple visual signals, including a conspicuous visual signal known as the foot flag, which is produced by rotating the hind limb above the body. However, FLUT treatment caused no measurable change in vocal signaling behavior, or in the frequency or fine temporal properties of males calls. Our study, therefore, suggests that activation of AR is not a physiological prerequisite to the coordination of multiple signals, in that it either does not regulate all signaling behaviors in a male s display repertoire or it does so only in a context-dependent manner

A Common Endocrine Signature Marks the Convergent Evolution of an Elaborate Dance Display in Frogs

Unrelated species often evolve similar phenotypic solutions to the same environmental problem, a phenomenon known as convergent evolution. But how do these common traits arise? We address this question from a physiological perspective by assessing how convergence of an elaborate gestural display in frogs (foot-flagging) is linked to changes in the androgenic hormone systems that underlie it. We show that the emergence of this rare display in unrelated anuran taxa is marked by a robust increase in the expression of androgen receptor (AR) messenger RNA in the musculature that actuates leg and foot movements, but we find no evidence of changes in the abundance of AR expression in these frogs’ central nervous systems. Meanwhile, the magnitude of the evolutionary change in muscular AR and its association with the origin of foot-flagging differ among clades, suggesting that these variables evolve together in a mosaic fashion. Finally, while gestural displays do differ between species, variation in the complexity of a foot-flagging routine does not predict differences in muscular AR. Altogether, these findings suggest that androgen-muscle interactions provide a conduit for convergence in sexual display behavior, potentially providing a path of least resistance for the evolution of motor performance

Multimodal Communication in a Noisy Environment: A Case Study of the Bornean Rock Frog Staurois parvus

High background noise is an impediment to signal detection and perception. We report the use of multiple solutions to improve signal perception in the acoustic and visual modality by the Bornean rock frog, Staurois parvus. We discovered that vocal communication was not impaired by continuous abiotic background noise characterised by fast-flowing water. Males modified amplitude, pitch, repetition rate and duration of notes within their advertisement call. The difference in sound pressure between advertisement calls and background noise at the call dominant frequency of 5578 Hz was 8 dB, a difference sufficient for receiver detection. In addition, males used several visual signals to communicate with conspecifics with foot flagging and foot flashing being the most common and conspicuous visual displays, followed by arm waving, upright posture, crouching, and an open-mouth display. We used acoustic playback experiments to test the efficacy-based alerting signal hypothesis of multimodal communication. In support of the alerting hypothesis, we found that acoustic signals and foot flagging are functionally linked with advertisement calling preceding foot flagging. We conclude that S. parvus has solved the problem of continuous broadband low-frequency noise by both modifying its advertisement call in multiple ways and by using numerous visual signals. This is the first example of a frog using multiple acoustic and visual solutions to communicate in an environment characterised by continuous noise

Divergent receiver responses to components of multimodal signals in two foot-flagging frog species.

Multimodal communication of acoustic and visual signals serves a vital role in the mating system of anuran amphibians. To understand signal evolution and function in multimodal signal design it is critical to test receiver responses to unimodal signal components versus multimodal composite signals. We investigated two anuran species displaying a conspicuous foot-flagging behavior in addition to or in combination with advertisement calls while announcing their signaling sites to conspecifics. To investigate the conspicuousness of the foot-flagging signals, we measured and compared spectral reflectance of foot webbings of Micrixalus saxicola and Staurois parvus using a spectrophotometer. We performed behavioral field experiments using a model frog including an extendable leg combined with acoustic playbacks to test receiver responses to acoustic, visual and combined audio-visual stimuli. Our results indicated that the foot webbings of S. parvus achieved a 13 times higher contrast against their visual background than feet of M. saxicola. The main response to all experimental stimuli in S. parvus was foot flagging, whereas M. saxicola responded primarily with calls but never foot flagged. Together these across-species differences suggest that in S. parvus foot-flagging behavior is applied as a salient and frequently used communicative signal during agonistic behavior, whereas we propose it constitutes an evolutionary nascent state in ritualization of the current fighting behavior in M. saxicola