The Function of Howling in the Ring-Tailed Lemur (Lemur catta)
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Long calls are sex-specific vocalizations used for mate attraction or mate defense in many animal species. Ring-tailed lemurs (Lemur catta), female-dominant strepsirrhines, have a male-specific long call termed a howl, with proposed functions that have never been empirically tested. I aimed to investigate why ring-tailed lemur males howl and to test whether the mate defense and mate attraction hypotheses for long-calling were applicable to this species. From March to July 2010, I collected 600 h of focal data on 25 males aged ≥3 year at Beza Mahafaly Special Reserve, Madagascar. I observed each male continuously for 30 min at a time and noted all agonism using one–zero sampling at 2.5-min intervals. I calculated male dominance rank from these data. I recorded days when female estrus occurred and noted howling and intergroup encounters using all-occurrences sampling. Howling rate was not significantly related to female estrus or male dominance rank, providing no support for the mate attraction hypothesis or the intragroup mate defense hypothesis. In contrast, the intergroup mate defense hypothesis was strongly supported. During intergroup encounters, male howling rate significantly increased compared to howling rate at times without other groups present, and a greater number of males participated in multimale howling choruses when compared to times without nongroup members present. My results suggest that male ring-tailed lemurs howl to advertise their presence and location to other groups, but not to male or female members of their own group. Howling could discourage male immigration by advertising the number of males already present in a group. Long calls are used for similar mate defense purposes during intergroup encounters by other primates, including Thomas langurs (Presbytis thomasi) and chacma baboons (Papio ursinus).
KeywordsLong call Loud call Mate attraction hypothesis Mate defense hypothesis
All data were collected in Madagascar with the approval of Madagascar National Parks (MNP), formerly known as Association Nationale pour la Gestion des Aires Protégées (ANGAP). I thank Jacky Ibrahim Antho Youssouf and Andry Randrianandrasana at the Beza Mahafaly Special Reserve for their facilitation of my project. My appreciation goes to Jeannin Rainavanosy and Joel Ratsirarson of the Département des Eaux et Forêts de l’Ecole Supérieure des Sciences Agronomiques (ESSA) and MNP/ANGAP for permission to work at Beza Mahafaly, and to Madagascar Institut pour la Conservation des Ecosystèmes Tropicaux (MICET) and Benjamin Andriamihaja for their help with logistics. I thank Elahavelo, Efitiria, Enafa, Edouard, Ralaevo, and Monja of the Beza Mahafaly Ecological Monitoring team for field assistance, and Teague O’Mara and his assistants for their help with orientation at Beza.
I sincerely thank Michelle Sauther and Frank Cuozzo for sharing ring-tailed lemur age data. I also thank Joyce Parga and Shawn Lehman for their excellent comments on this manuscript, Drew Rendall and Shannon Digweed for introducing me to acoustic research, and Ryan Janzen for technological aid. Finally, I am grateful to Joanna Setchell and two anonymous reviewers for their helpful suggestions. For funding, I acknowledge an Alexander Graham Bell Canada Graduate Scholarship from the Natural Sciences and Engineering Research Council of Canada, the American Museum of Natural History, the Edward. J. Noble Foundation, St. Catherines Island Foundation, Ontario Graduate Scholarships, and the University of Toronto.
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