Sexual signalling in Propithecus verreauxi: male "chest badge" and female mate choice.

Communication, an essential prerequisite for sociality, involves the transmission of signals. A signal can be defined as any action or trait produced by one animal, the sender, that produces a change in the behaviour of another animal, the receiver. Secondary sexual signals are often used for mate choice because they may inform on a potential partner's quality. Verreaux's sifaka (Propithecus verreauxi) is characterized by the presence of two different morphs of males (bimorphism), which can show either a stained or clean chest. The chest becomes stained by secretions of the sternal gland during throat marking (rubbing throat and chest on a vertical substrate while smearing the scent deposition). The role of the chest staining in guiding female mate choice was previously hypothesized but never demonstrated probably due to the difficulty of observing sifaka copulations in the wild. Here we report that stained-chested males had a higher throat marking activity than clean-chested males during the mating season, but not during the birth season. We found that females copulated more frequently with stained-chested males than the clean-chested males. Finally, in agreement with the biological market theory, we found that clean-chested males, with a lower scent-releasing potential, offered more grooming to females. This "grooming for sex" tactic was not completely unsuccessful; in fact, half of the clean-chested males copulated with females, even though at low frequency. In conclusion, the chest stain, possibly correlated with different cues targeted by females, could be one of the parameters which help females in selecting mates.

Scratching around mating: factors affecting anxiety in wild Lemur catta.

Scratching has been successfully used to detect anxiety, a proxy for stress, in primates, from strepsirrhines to Homo sapiens. Here, we investigated the fluctuation of scratching in Lemur catta during the mating season. In particular we evaluated whether scratching (1) varied according to sex and rank differences, (2) increased in the period of maximum stress (around the mating days), and (3) was reduced by grooming. At Berenty (South Madagascar), we followed two lemur groups (23 adult/subadult individuals) and gathered data on self-scratching, aggression, and grooming. Based on perineal area features, we recognized two periods: low swelling (LS), with no estrus female, and high swelling (HS), when at least one female was in estrus. We predicted that aggressive behaviors and anxiety-related scratching would covary. Indeed, scratching peaked in HS, when aggression was also highest. In agreement with previous literature, this result suggests that conflicts around estrus days may raise anxiety levels in the social group. We expected scratching levels to be highest in males because they aggressively compete for females and are subject to mate choice and repeated attacks by dominant females. Instead, the scratching rates were similar in males and females, probably because the high competition, which involves both sexes, dampened intersexual differences. In contrast to our prediction, scratching was not rank dependent, probably because animal ranking positions changed from LS to HS. Finally, we showed that, in ring-tailed lemurs, as well as in other primates, scratching decreases after reciprocal grooming in both periods. This finding provides the first evidence that grooming could assist in reducing anxiety in strepsirrhines.

Stranger to familiar: wild strepsirhines manage xenophobia by playing.

The power of play in limiting xenophobia is a well-known phenomenon in humans. Yet, the evidence in social animals remains meager. Here, we aim to determine whether play promotes social tolerance toward strangers in one of the most basal group of primates, the strepsirhines. We observed two groups of wild lemurs (Propithecus verreauxi, Verreaux's sifaka) during the mating season. Data were also collected on nine visiting, outgroup males. We compared the distribution of play, grooming, and aggressive interactions across three conditions: OUT (resident/outgroup interactions), IN (resident/resident interactions in presence of outgroups) and BL-IN (baseline of resident/resident interactions in absence of outgroups). Play frequency between males was higher in OUT than in IN and BL-IN conditions; whereas, grooming was more frequent in IN than in OUT and BL-IN conditions. Aggression rates between resident and outgroup males were significantly higher than those between residents. However, aggressions between resident and outgroup males significantly decreased after the first play session and became comparable with resident-resident aggression levels. The presence of strangers in a well-established group implies the onset of novel social circumstances, which sifaka males cope with by two different tactics: grooming with ingroup males and playing with outgroup ones. The grooming peak, concurrently with the visit of outgroups, probably represents a social shield adopted by resident males to make their pre-existing affiliation more evident to the stranger "audience". Being mostly restricted to unfamiliar males, adult play in sifaka appears to have a role in managing new social situations more than in maintaining old relationships. In particular, our results indicate not only that play is the interface between strangers but also that it has a specific function in reducing xenophobia. In conclusion, play appears to be an ice-breaker mechanism in the critical process that "upgrades" an individual from stranger to familiar.

Mating first, mating more: biological market fluctuation in a wild prosimian.

In biology, economics, and politics, distributive power is the key for understanding asymmetrical relationships and it can be obtained by force (dominance) or trading (leverage). Whenever males cannot use force, they largely depend on females for breeding opportunities and the balance of power tilts in favour of females. Thus, males are expected not only to compete within their sex-class but also to exchange services with the opposite sex. Does this mating market, described for humans and apes, apply also to prosimians, the most ancestral primate group? To answer the question, we studied a scent-oriented and gregarious lemur, Propithecus verreauxi (sifaka), showing female dominance, promiscuous mating, and seasonal breeding. We collected 57 copulations involving 8 males and 4 females in the wild (Berenty Reserve, South Madagascar), and data (all occurrences) on grooming, aggressions, and marking behaviour. We performed the analyses via exact Spearman and matrix correlations. Male mating priority rank correlated with the frequency of male countermarking over female scents but not with the proportion of fights won by males over females. Thus, males competed in an olfactory tournament more than in an arena of aggressive encounters. The copulation frequency correlated neither with the proportion of fights won by males nor with the frequency of male countermarking on female scents. Male-to-female grooming correlated with female-to-male grooming only during premating. Instead, in the mating period male-to-female grooming correlated with the copulation frequency. In short, the biological market underwent seasonal fluctuations, since males bargained grooming for sex in the mating days and grooming for itself in the premating period. Top scent-releasers gained mating priority (they mated first) and top groomers ensured a higher number of renewed copulations (they mated more). In conclusion, males maximize their reproduction probability by adopting a double tactic and by following market fluctuations.

Fine-tuning of social play in juvenile lowland gorillas (gorilla gorilla gorilla).

Social play, which involves cooperation, communication, and learning, may represent a suitable field for the investigation of cognitive ability in a given species. We collected data on a captive group of gorillas in order to evaluate the potential cognitive skill of juveniles in fine-tuning play behavior. This study revealed that juvenile gorillas are able to "place" the play session in a proper spatial/temporal context, thus evaluating a complex net of factors (e.g., play partner, play roughness, group activity, space availability). When animals play fight, they use patterns of agonistic functional contexts. Since these actions are not intrinsically different from their "serious" context, it may be hard to distinguish them. One of the most important function of play in the ontogeny of primate social cognition may be to recognize stimuli, which may indicate the intentions of conspecifics. Accordingly, we found that juvenile gorillas are able to use play signals appropriately when a clear statement of purpose is necessary (i.e., during male-male competitive play sessions and when the escape opportunities are limited). The ability to interpret such ambiguous features of social signaling could represent a central issue in the evolution of behavioral flexibility and intelligence in primates.