Animal communication signals can be highly elaborate, and researchers have long sought explanations for their evolutionary origins. For example, how did signals such as the tail-fan display of a peacock, a firefly flash or a wolf howl evolve? Animal communication theory holds that many signals evolved from non-signalling behaviours through the process of ritualization. Empirical evidence for ritualization is limited, as it is necessary to examine living relatives with varying degrees of signal evolution within a phylogenetic framework. We examine the origins of vibratory territorial signals in caterpillars using comparative and molecular phylogenetic methods. We show that a highly ritualized vibratory signal—anal scraping—originated from a locomotory behaviour—walking. Furthermore, comparative behavioural analysis supports the hypothesis that ritualized vibratory signals derive from physical fighting behaviours. Thus, contestants signal their opponents to avoid the cost of fighting. Our study provides experimental evidence for the origins of a complex communication signal, through the process of ritualization.

Vibrational communication in hook-tip moth caterpillars is thought to be widely used and highly variable across species, but this phenomenon has been experimentally examined in only two species to date. The purpose of this study is to characterize and describe the function of vibrational signaling in a species, Oreta rosea Walker 1855 (Lepidoptera: Drepanidae), that differs morphologically from previously studied species. Caterpillars of this species produce three distinct types of vibrational signals during territorial encounters with conspecifics — mandible drumming, mandible scraping and lateral tremulation. Signals were recorded using a laser-doppler vibrometer and characterized based on temporal and spectral components. Behavioural encounters between a leaf resident and a conspecific intruder were staged to test the hypothesis that signaling functions as a territorial display. Drumming and scraping signals both involve the use of the mandibles, being hit vertically on, or scraped laterally across, the leaf surface. Lateral tremulation involves quick, short, successive lateral movements of the anterior body region that vibrates the entire leaf. Encounters result in residents signaling, with the highest rates observed when intruders make contact with the resident. Residents signal significantly more than intruders and most conflicts are resolved within 10 minutes, with residents winning 91% of trials. The results support the hypothesis that vibrational signals function to advertise leaf occupancy. Signaling is compared between species, and evolutionary origins of vibrational communication in caterpillars are discussed.

Abstract. Caterpillars of the poplar lutestring moth, Tethea or, construct leaf shelters that they defend against intruding conspecifics using a combination of vibratory signals and physical aggression. Staged interactions between a resident caterpillar and introduced conspecific were recorded with a video camera and laser vibrometer. Residents crawl towards the intruder and perform three behaviours: lateral hitting, pushing, and mandible scraping. Vibrations caused by mandible scraping result from the caterpillar repeatedly scraping opened mandibles laterally against the leaf surface in bouts lasting 1.16 ± 0.39 s, with an average of 4 ± 1 scrapes per bout. We propose that these scrapes function in leaf shelter defense against conspecifics for the following reasons: Mandible scrapes are produced only by residents; they are generated when a resident is approached by an intruder; the rate of scraping increases as the intruder approaches the shelter; and residents in all trials retain their shelters, with the intruder leaving the leaf within 127.9 ± 104.3 s from the beginning of the trial. The function and evolutionary origins of vibration-mediated territoriality in caterpillars are discussed.