Primates are traditionally considered to be microsmatic, with decreased reliance on olfactory senses in comparison to other sensory modalities such as vision. This is particularly the case for Old World monkeys and apes (catarrhines). However, various lines of evidence suggest that chemical communication may be important in these species, including the presence of a sternal scent-gland in the mandrill. We investigated the volatile components of mandrill odor using gas chromatography-mass spectrometry. We identified a total of 97 volatile components in 88 swabs of the sternal gland secretion and 95 samples of sternal gland hair saturated with scent-gland secretion collected from 27 males and 18 females. We compared odor profiles with features of the signaler using principle components and discriminant function analyses and found that volatile profiles convey both variable (age, dominance rank in males) and fixed (sex, possibly individual identity) information about the signaler. The combination of an odor profile that signals sex, age, and rank with increased motivation to scent-mark and increased production of secretion in high-ranking males leads to a potent signal of the presence of a dominant, adult male with high testosterone levels. This may be particularly relevant in the dense Central African rain forest which mandrills inhabit. By contrast, we were unable to differentiate between either female cycle stage or female rank based on odor profiles, which accords with behavioral studies suggesting that odor signals are not as important in female mandrills as they are in males. The similarity of our findings to those for other mammals and in primates that are more distantly related to humans suggests a broader role for odor in primate communication than is currently recognized.

The major histocompatibility complex (MHC) is an extraordinarily diverse cluster of genes that play a key role in the immune system. MHC gene products are also found in various body secretions, leading to the suggestion that MHC genotypes are linked to unique individual odourtypes that animals use to assess the suitability of other individuals as potential mates or social partners. We investigated the relationship between chemical odour profiles and genotype in a large, naturally reproducing population of mandrills, using gas chromatography-mass spectrometry and MHC genotyping. Odour profiles were not linked to the possession of particular MHC supertypes. Sex influenced some measures of odour diversity and dominance rank influenced some measures of odour diversity in males, but not in females. Odour similarity was strongly related to similarity at the MHC, and, in some cases, to pedigree relatedness. Our results suggest that odour provides both a cue of individual genetic quality and information against which the receiver can compare its own genotype to assess genetic similarity. These findings provide a potential mechanism underlying mate choice for genetic diversity and MHC similarity as well as kin selection.

Mandrills are one of the few Old World primates to show scent-marking. We combined ethological and chemical approaches to improve our understanding of this behavior in 3 zoo-managed groups. We observed the olfactory behavior performed by adults and adolescents (N = 39) for 775h. We investigated the volatile components of sternal scent-marks using gas chromatography-mass spectrometry and compared volatile profiles with traits of the signaler. Males marked more than females and within each sex the frequency of scent-marking was related to age and dominance status, but alpha males scent-marked most frequently and particularly in specific areas at the enclosure boundaries. We identified a total of 77 volatile components of sternal gland secretion, including compounds functioning as male sex pheromones in other mammals, in scent-marks spontaneously released on filter paper by 27 male and 18 female mandrills. We confirmed our previous findings that chemical profiles contain information including sex, male age and rank, and we also found that odor may encode information about group membership in mandrills. Our results support the hypotheses that scent-marking signals the status of the dominant male as well as playing territorial functions but also suggest that it is part of sociosexual communication.

Primate chemical communication remains underappreciated, as primates are considered to rely on other sensory modalities. However, various lines of evidence suggest that olfaction plays an important role in primate societies, including the conspicuous scent-marking behavior of many strepsirrhines and callitrichines. Although lemurs typically show scent-marking, little is known about this behavior in red-ruffed lemurs (Varecia variegata rubra). We combined behavioral observations and semiochemistry analyses to improve our understanding of scent-marking in two captive troops housed at Dudley and Twycross zoos (UK). We collected olfactory behavioral observations by focusing on two family troops (N = 7) for 132 hr. We investigated the volatile compounds of ano-genital scent-marks using solid-phase microextraction and gas chromatography-mass spectrometry and compared volatile chemical profiles with features of the signaller. Males scent-marked most frequently and predominantly in specific meaningful areas of the enclosure, while within females the occurrence of scent-marking was related to their age. We found behavioral sexual dimorphism, with male predominantly depositing secretions via neck and mandible glands and females via ano-genital glands. We identified a total of 32 volatile components of ano-genital gland secretion, including compounds that have already been found in other mammals as sex pheromones and cues to fitness, in ano-genital scent-marks spontaneously left on filter paper by adult females. Our findings suggest that red-ruffed lemurs might use scent-marking to convey information about sex and female age, with male neckmarking behavior playing defensive territorial functions and ano-genital marking related to socio-sexual communication. K E Y W O R D S communication, gas chromatography-mass spectrometry, olfaction, signaling, Varecia variegata rubra

Female primates signal impending ovulation with a suite of sexual signals. Studies of these signals have focussed on visual, and to a lesser extent, acoustic signals, neglecting olfactory signals. We aimed to investigate the information content of female olfactory signals in captive olive baboons (Papio anubis) and relate these to the female fertile period. We studied eight adult females living in four groups at the CNRS Station de Primatologie, Rousset-sur-Arc, France. We used vaginal cytology to detect ovulation. We investigated the volatile component of odour signals using solid-phase microextraction and gas chromatography-mass spectrometry. We found a total of 74 volatile compounds, of which we tentatively identified 25, including several ketones, alcohols, aldehydes, terpenes, volatile fatty acids and hydrocarbons that have been identified in odour profiles of other primates. Our results show that vaginal odour intensity differs with sexual cycle stage suggesting that odour might play a role in signalling female baboon fertility. We found differences in vaginal odour between females living in all-female and in mixed sex groups but we could not distinguish the effects of group composition, female age and identity. This study of olfactory signalling improves our understanding of how female primates advertise their sexual receptivity.

Publisher's copyright statement:The nal publication is available at Springer via http://dx.doi.org/10.1007/s10886-008-9573-5. Use policyThe full-text may be used and/or reproduced, and given to third parties in any format or medium, without prior permission or charge, for personal research or study, educational, or not-for-prot purposes provided that:• a full bibliographic reference is made to the original source • a link is made to the metadata record in DRO • the full-text is not changed in any way The full-text must not be sold in any format or medium without the formal permission of the copyright holders.Please consult the full DRO policy for further details. Abstract Human pheromones have a role in regulating relationships, and apparently influence partner choice and mother-infant recognition. We analyzed the chemical content of volatiles from sweat patch samples from the para-axillary and nipple-areola regions of women during pregnancy and after childbirth. Solid phase microextraction (SPME) was used to extract the volatile compounds, which were then characterized and quantified by gas chromatography-mass spectrometry (GC-MS). During pregnancy, women developed a distinctive pattern of five volatile compounds common to the para-axillary and nipple-areola regions (1-dodecanol, 1-1΄-oxybis octane, isocurcumenol, α-hexyl-cinnamic aldehyde, and isopropyl myristate). These compounds were absent outside pregnancy and had slightly different patterns in samples from the two body areas. Differentiation of the volatile patterns among pregnant women may help newborns to distinguish their own mothers.

Positive reinforcement training (PRT) is an established tool to facilitate animal husbandry, care and research in modern zoos, with potential positive implications for captive animal welfare. The study explored the role of an isolation PRT training programme on the well-being of ring-tailed lemurs (Lemur catta). Eleven subjects were observed during an isolation training protocol to induce the animals to enter an area (training area) calmly and retrieve rewards separated from group members. Duration of individual and social behaviours were collected over two different periods: the baseline period, before the beginning of the isolation training protocol and the training period, in which the collection of the data started at the end of the isolation training sessions. Additionally, behavioural data within the isolation training sessions (latency to enter the training area and retrieve the reward, display of stress-related behaviours) were recorded. Outside the training sessions, lemurs were out of sight significantly more in the baseline (Mean ± SD: 15.46 ± 5.20) than in the training (Mean ± SD: 4.36 ± 2.89) period. Social behaviour was performed significantly more in the training (Mean ± SD: 31.80 ± 12.34) than in the baseline (Mean ± SD: 12.52 ± 5.14) period; particularly, lemurs were in social contact significantly more in the training (Mean ± SD: 14.09 ± 6.00) than in the baseline period (Mean ± SD: 4.58 ± 2.73). Agonistic behaviours were performed significantly more in the baseline (Mean ± SD: 0.23 ± 0.15) than in the training (Mean ± SD: 0.07 ± 0.07) period. Within the training sessions, all the individuals entered the training area, were isolated from conspecifics, and retrieved the reward in 6 out of 9 sessions. Our findings show that, during the PRT period, lemurs displayed their natural behaviour in their everyday social life with significant increase of their affiliative behaviours and decrease of aggressive behaviours with benefits for their welfare status. Thus, lemurs were able to cope with the use of PRT to isolate each individual from its social group − a situation which, without training, might be very stressful. In conclusion, PRT may play a crucial role for the captive management of ring-tailed lemurs in captive facilities, including zoos

Fifty years after the term "pheromone" was coined by Peter Karlson and Martin Lüsher the search for these semiochemicals is still an elusive goal of chemical ecology and communication studies. Contrary to what appears in the popular press, the race is still on to capture and define human scents. Over the last several years, it became increasingly clear that pheromone-like chemical signals probably play a role in offspring identification and mother recognition. Recently, we analyzed the volatile compounds in sweat patch samples collected from the para-axillary and nipple-areola regions of women during pregnancy and after childbirth. We hypothesized that, at the time of birth and during the first weeks of life, the distinctive olfactory pattern of the para-axillary area is probably useful to newborns for recognizing and distinguishing their own mother, whereas the characteristic pattern of the nipple-areola region is probably useful as a guide to nourishment.