Traditionally, frogs and toads are considered to communicate primarily by using acoustic cues, with males typically calling to attract females. However, in noisy tropical rainforests by fast flowing streams, acoustic communication becomes more problematic. In these environments male frogs have evolved an array of visual cues which complement acoustic cues to communicate between each other and to also attract females. Visual communication has evolved independently in several lineages of frog species, mainly in diurnal species inhabiting noisy, stream-side environments.
Foot-flagging has been observed in 16 frog species (Preininger et al., 2013). The male will typically arch and rotate its back foot in the air, giving a conspicuous visual signal (Figure 1). Males of the Bornean frog genus Staurois have evolved a unique foot-flagging behaviour which males use in a variety of contexts. The Sabah splash frog (Staurois latopalmatus) is found exclusively close to waterfalls where it can be observed regularly on exposed perches. Within this habitat, males signal in close vicinity to each other near to the water (Preininger et al., 2009). Preininger et al. (2009) have identified three types of visual displays: foot flagging, arm waving and vocal sac displays. Foot-flagging displays are mainly performed in the direction of a male opponent and appear to be used in territorial defence. Preininger et al. (2009) hypothesise that foot-flagging behaviour in this species has evolved from physical male-male combat where each male tries to push the other off its perch. The foot-flagging behaviour may be a ritualization of this aggressive combat, avoiding the need for physical contact. The other visual displays, arm waving and throat display appear to be used less frequently than foot-flagging during male-male aggressive encounters.
The Bornean rock frog (Staurois parvus) from Borneo and the small torrent frog (Micrixalus saxicola) from the Western Ghats of India belong to different frog families (Figure 2). Males of both species use complex signalling involving high pitched calls, foot flagging, and tapping (foot lifting) to defend perching sites against other males (Preininger et al. 2013). The Bornean rock frog has conspicuous white feet, whereas the small torrent frog has feet which are the same colour as its body. In a study to examine the differences in the behaviour of the two species, Preininger et al. (2013) found that in the Bornean rock frog, foot-flagging achieved a 13 times higher contrast against their visual background than the feet of the small torrent frog. In addition, the Bornean rock frog primarily responded to stimuli with foot flagging, whereas the small torrent frog responded mainly with calls but never foot-flagging on its own (Preininger et al. 2013). The authors propose that in the small torrent frog foot-flagging is in a transient state, evolving from its current use in physical fighting behaviour.
The colouration on the feet of foot-flagging species may signal more than male presence. Research by Stangel et al. (2015) has found that in two species (Staurois parvus and S. guttatus) the brightness of the feet increases with age. The peak brightness seems to coincide with sexual maturity and may be linked to androgen hormone levels in the males. The foot-flagging in these species may therefore convey information on the status of the male and his receptiveness to mate. This may be useful both to intruding males and also females which may be in the area.
African puddle frogs of the genus Phrynobatrachus are unique to Africa and are named after their breeding strategy of often laying large numbers of eggs in slow-moving or stagnant water bodies. In East African mountain streams lives a day time active frog, krefft’s river frog (Phrynobatrachus krefftii). Like other members of its genus males are dull brown in colour but also possess a striking bright yellow vocal sac (Figure 3). Instead of using foot-flagging for visual communication, males of this species communicate using a combination of acoustic cues and exhibiting their bright yellow vocal sac. Sometimes males will utter a call, accompanied by exhibiting their yellow vocal sac, whereas at other times no call will be emitted. Hirschmann & Hödl (2006) propose that the use of their brightly coloured vocal sac in this way has evolved to indicate the aggressive motivational state in the male.
Further research into foot-flagging and other visual displays at different life stages will help increase our understanding of the function, development and evolution of visual signals in amphibians. Foot-flagging is only confined to a relatively few number of frog species and these are unable to survive in habitats modified for human use. Habitats where many of these unique species occur are increasingly threatened with habitat loss and fragmentation. Understanding the ecology of these species is therefore crucial in identifying and protecting key habitats in the wild.
Amézquita, A. & Hödl, W. (2004) How, when and where to perform visual displays: the case of the Amazonian frog Hyla parviceps. Herpetologica, 60 (4): 420–429.
Hirschmann, W. & Hödl, W. (2006) Visual signaling in Phrynobatrachus krefftii Boulenger, 1909 (Anura: Ranidae). Herpetologica, 62 (1): 18–27.
Preininger, D., Boechle, M. & Hödl, W. (2009) Communication in noisy environments II: visual signaling behavior of male foot-flagging frogs Staurois latopalmatus. Herpetologica, 65 (2): 166–173.
Preininger, D., Boechle, M., Sztatecsny, M. & Hödl, W. (2013) Divergent receiver responses to components of multimodal signals in two foot-flagging frog species. PLoS ONE, 8 (1): e55367. doi:10.1371/journal.pone.0055367.
Stangel, J., Preininger, D., Sztatecsny, M. & Hödl, W. (2015) Ontogenetic change of signal brightness in the foot-flagging frog species Staurois parvus and Staurois guttatus. Herpetologica, 71 (1): 1–7.