Mantled howler monkey males assess their rivals through formant spacing of long-distance calls

Formant frequency spacing of long-distance vocalizations is allometrically related to body size and could represent an honest signal of fighting potential. There is, however, only limited evidence that primates use formant spacing to assess the competitive potential of rivals during interactions with extragroup males, a risky context. We hypothesized that if formant spacing of long-distance calls is inversely related to the fighting potential of male mantled howler monkeys (Alouatta palliata), then males should: (1) be more likely and (2) faster to display vocal responses to calling rivals; (3) be more likely and (4) faster to approach calling rivals; and have higher fecal (5) glucocorticoid and (6) testosterone metabolite concentrations in response to rivals calling at intermediate and high formant spacing than to those with low formant spacing. We studied the behavioral responses of 11 adult males to playback experiments of long-distance calls from unknown individuals with low (i.e., emulating large individuals), intermediate, and high (i.e., small individuals) formant spacing (n = 36 experiments). We assayed fecal glucocorticoid and testosterone metabolite concentrations (n = 174). Playbacks always elicited vocal responses, but males responded quicker to intermediate than to low formant spacing playbacks. Low formant spacing calls were less likely to elicit approaches whereas high formant spacing calls resulted in quicker approaches. Males showed stronger hormonal responses to low than to both intermediate and high formant spacing calls. It is possible that males do not escalate conflicts with rivals with low formant spacing calls if these are perceived as large, and against whom winning probabilities should decrease and confrontation costs increase; but are willing to escalate conflicts with rivals of high formant spacing. Formant spacing may therefore be an important signal for rival assessment in this species.