Defensive and Exploratory Reactions of Black and Golden-Headed Lion Tamarins Exposed to a Mirror
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Revista de Etologia 2001, Vol.3, N°1, 15-23 Defensive and exploratory responses to mirrors in lion tamarins The Image Is Another Tamarin: Defensive and Exploratory Reactions of Black and Golden-Headed Lion Tamarins Exposed to a Mirror CÉSAR ADES, LUCIANO TOMÉ AND VÂNIA HADDAD DIEGO Universidade de São Paulo To determine the nature of the social behavior of black (Leontopithecus chrysopygus) and golden-faced (L. chrysomelas) lion tamarins toward their mirror-image, the responses of lion tamarins of both species (defense, exploration, observation of mirror and stay in proximity of the mirror) were assessed in their home cages during two sessions of exposure to mirrors and compared to baseline levels. Mirrors elicited, on first and second presentation, strong defensive responses such as vocalizations, scent marking, increased locomotion, as well as exploration and observation directed to the mirror. On the second exposure, there was a decrease in defensive performance for black but not for golden-headed lion tamarins, while exploratory and looking to the mirror levels remained at their high, former levels. Results indicate that lion tamarins, contrary to some indication of non-aggressive responsiveness to mirror-image in cotton-top tamarins, reacts defensively to mirror-images as though they were interacting with an intruder. Permanence of observing responses to the mirror-image, at least in black lion tamarins, may indicate the existence of a social exploration tendency, independent from aggressiveness toward strange conspecifics. Index terms: Mirrors, Defensive behavior. Lion tamarins. Leontopithecus chrysopygus, L. chrysomelas. A imagem é outro mico: reações defensivas e exploratórias de micos-leões-pretos e micos-leões-de-cara- dourada diante do espelho. A fim de determinar a natureza do comportamento social de micos-leões-pretos (Leontopithecus chrysopygus) e micos-leões-de-cara-dourada (L. chrysomelas) diante de sua imagem ao espelho, fo- ram registradas por duas sessões, nos seus recintos, as respostas ao espelho de indivíduos de ambas as espécies – defesa, exploração, olhar para o espelho e permanência próximo ao espelho – comparando os níveis encontra- dos aos da linha de base. O espelho provocou respostas defensivas intensas – vocalizações, marcação de cheiro, locomoção, etc. - em ambas as apresentações, assim como comportamentos de exploração e de observação dirigidos ao espelho. Os micos-leões-pretos, mas não os micos-leões-de-cara-dourada, diminuiram seu compor- tamento defensivo na segunda sessão, mantendo contudo altos a exploração e a observação do espelho. Os resultados indicam que, contrariamente a relatos sobre a pouca agressividade de sagüis-de-topete diante do espelho, os micos-leões reagem defensivamente à sua imagem, como se ela representasse um indivíduo estra- nho de sua espécie. A permanência das respostas de observação do espelho, pelo menos em micos-leões-pretos, pode ser uma indicação da existência de uma tendência para a exploração social, independente da agressividade. Descritores: Espelhos. Comportamento defensivo. Micos-leões. Leontopithecus chrysopygus. L. chrysomelas. Many animals ignore their image seen in (MacLean, 1964), pygmy marmosets Cebuella a mirror, but in some species with well- pygmea (Eglash & Snowdon, 1983) and Cerra- developed vision, particularly primates, mirrors may elicitate species-typical exploratory or so- Thanks are due to the chrysopygus (Elisabete C. Lopes, cial responses. Among the New World monkeys, Karin A. Onodera e Úrsula M. Liemert) and to the chrysomelas (Mariana M. Kroll, Lucas R. Napolitano, capuchins Cebus apella (Anderson & Roeder, Gigliana S. Lima, Liana L. de Souza and Marcos Takahashi) 1989) squirrel monkeys, Saimiri sciureus teams for cooperation and enthusiasm during data collection, to the Fundação Parque Zoológico de São Pau- César Ades, Instituto de Psicologia, USP. Av. Prof. Mello lo for the permission to perform our experiment with lion Moraes, 1721, 05508-900. E-mail:[email protected] tamarins and to FAPESP and CNPq for research grants. 15 César Ades, Luciano Tomé and Vânia Haddad Diego do marmosets Callithrix penicillata (Perin & Guer- touching the place on their head where hair had ra, 2000) have been observed to react with been dyed. Again, tamarins “stared into the increased curiosity and/or aggressiveness to mirror peacefully and for long durations, but their mirror-image, at least during the first did not vocalize or act aggressively” (p. 10813). exposures, in a manner which suggest that they They did so for longer periods than they had may take the image for a conspecific. done before their hair changed colors. Hauser, Mirror tests have also been used as a Miller, Liu, and Gupta (2001) recently technique for assessing self-recognition in performed a replication of Hauser et al. (1995) primates. Chimpanzees may react to their image experiment, as well as a new one designed to with threatening displays but on repeated follow closely the technique used with exposure, they engage in a variety of self- chimpanzees, and failed, in both, to find any directed behaviors when looking at themselves evidence of self-recognition as indicated by (such as watching inside their own mouth), and touches to the dye-marked area. Animals may inspect and touch parts of their face however tended to look longer at the mirror – marked with dye which they detect through the relatively to habituation sessions – when first mirror-image (Gallup, 1977; Povinelli, Rulf, exposed to their new, colored visual Landau, & Bierschwale, 1993). Self directed appearance. Hauser et al. (2001) again mention behavior at the mirror, reported in chimpanzees that during familiarization to the mirror, in and orangutans but not in gorillas or in Experiment 2, no head touching and no monkeys, has been interpreted by Gallup (1998) aggressive displays toward the mirror were as indicating that, in some apes at least, a fun- recorded. During test sessions, animals did not damental evolutionary step towards self- direct prolonged looks to the mirror, nor did consciousness and towards attribution of they display aggressively. consciousness to conspecifics was taken (see This lack or low frequency of agonistic Ades, 1998; Pereira Jr., 1999, and de Veer e van responses, in both studies of Hauser et al. (1995, den Bos, 1999 for critical discussions of such 2001) is somewhat surprising once, in many interpretations). species of primates, mirror images elicit intense Our interest in testing lion tamarins in a and prolonged threat and defensive displays. mirror context was aroused by the experiment The purpose of the present research, performed on self-recognition Hauser, Kralik, Botto- with captive black (Leonthopithecus chrysopygus) Mahan, Garrett, and Oser (1995) conducted on and golden-headed lion tamarins (L. cotton-tops tamarins (Saguinus oedipus). Mirrors chrysomelas), was not to find an evidence of self- were mounted in the cages of tamarins for a knowledge through a Gallup mark test, but to period before experimental manipulations were investigate the nature of the reactions of lion performed. First contact with mirrors was tamarins to mirrors and to discover whether generally not aggressive. As Hauser et al. (1995) they are basically of the self or conspecific- put it: “tamarins with access to mirrors showed directed type. Our hypothesis was that the initial interest, evidenced by brief stares, visual monkeys would react with threat and other and manual searches behind the mirror, and defensive responses to their appearance in the occasional threats … mirror stares appeared mirror, as they would react to an intruder generally nonaggressive, with no accompanying conspecific. vocalizations and only five threat displays” (p. Lion tamarins, the larger of callithrichids, 10811). A mark test, analogous to the one are endangered species which live in swamp and performed on chimpanzees, was performed by dryland forests (Albernaz, 1997), in nuclear or dying the white hair on the tamarins’ head with extended families. While there may be ingroup bright colours. Only individuals with long aggressiveness, threats and aggressive categories previous exposures to the mirror were observed of behavior are mostly directed towards 16 Defensive and exploratory responses to mirrors in lion tamarins members of other groups. “Visual contact often with a front wire mesh partition, a small shelter leads to high arousal with considerable in its rear part and several tree trunks, two of locomotion, piloerection, and vocalizing. In our which served as bridges between shelter and the experience, injuries often result when two fore part of the enclosure. captive groups have had the opportunity to At the beginning of the sessions, a 50 x make physical contact through small openings 30 cm metallic, horizontal tray with a 50 x 55 between enclosures” (Kleiman, 1984, p. 39). cm vertical mirror above it, were mounted on Intergroup aggressive encounters in the field the wire mesh front partition, at approximately which have to do with feeding resource 1,5 m from the floor. Flour beetle larvae and competition involve visual contact, alarm calls, gum candies were spread on the tray, in all scent marking and sequences of chase and sessions, to attract tamarins. Animals were retreats, as in other callithrichid species. In his subjected to a 15-min baseline session with the research with wild golden lion tamarins L. rosalia mirror in a control position, opaque surface at the Poço das Antas Biological Reserve, Peres facing the enclosure,