Male Scent Marks Research Articles

Regulation of Sexually Dimorphic Expression of Major Urinary Proteins.

Male house mice excrete large amounts of protein in their urinary scent marks, mainly composed of Major Urinary Proteins (MUPs), and these lipocalins function as pheromones and pheromone carriers. Here, we review studies on sexually dimorphic MUP expression in house mice, including the proximate mechanisms controlling MUP gene expression and their adaptive functions. Males excrete 2 to 8 times more urinary protein than females, though there is enormous variation in gene expression across loci in both sexes. MUP expression is dynamically regulated depending upon a variety of factors. Males regulate MUP expression according to social status, whereas females do not, and males regulate expression depending upon health and condition. Male-biased MUP expression is regulated by pituitary secretion of growth hormone (GH), which binds receptors in the liver, activating the JAK2-STAT5 signaling pathway, chromatin accessibility, and MUP gene transcription. Pulsatile male GH secretion is feminized by several factors, including caloric restriction, microbiota depletion, and aging, which helps explain condition-dependent MUP expression. If MUP production has sex-specific fitness optima, then this should generate sexual antagonism over allelic expression (intra-locus sexual conflict) selectively favoring sexually dimorphic expression. MUPs influence the sexual attractiveness of male urinary odor and increased urinary protein excretion is correlated with the reproductive success of males but not females. This finding could explain the selective maintenance of sexually dimorphic MUP expression. Producing MUPs entails energetic costs, but increased excretion may reduce the net energetic costs and predation risks from male scent marking as well as prolong the release of chemical signals. MUPs may also provide physiological benefits, including regulating metabolic rate and toxin removal, which may have sex-specific effects on survival. A phylogenetic analysis on the origins of male-biased MUP gene expression in Mus musculus suggests that this sexual dimorphism evolved by increasing male MUP expression rather than reducing female expression.

Are male meadow voles (Microtus pennsylvanicus) influenced by social odor context with regard to scent marking behaviors?

AbstractMeadow voles (Microtus pennsylvanicus) rely on olfactory communication, such as scent marking, to find and secure mating opportunities during the breeding season. Scent marks, and odors in general, provide information about the donors and persist in the environment for extended periods, allowing a wide audience. Whether a vole responds to another scent mark(s), or odor(s), depends on the social context and the information the odor provides about the donor(s). Therefore, we examined if male scent marking toward a social odor or female meadow voles is influenced by information present or previously encountered based on contextual differences in the social odor. We presented males with two odor associations of a male with a female with contextual differences in the age of the rival. We tested male scent marking under two conditions: one with two social odors present, current information, and a second with the females present from previously encountered social information, prior information. It was found that rival male age did not affect the scent marking behavior of males toward a social odor or female voles. Rival male age also did not affect preference as males did not show a preference for either female. The lack of an effect may lie in the properties of scent marks and the natural history of meadow voles during the breeding season. The odor of an individual rival may not represent a strong enough competitive threat to influence a signaler's decisions, but the association could still provide the signaler with valuable information about which females are considered high quality.

How to maintain underground social relationships? Chemosensory sex, partner and self recognition in a fossorial amphisbaenian.

Maintaining social relationships depends on the ability to recognize partners or group members against other individuals. This is especially important in animals with relatively stable social groups. The amphisbaenian Trogonophis wiegmanni is a semi blind fossorial reptile that spends its entire life underground where it interacts with mates and social partners. In this environment, visual cues are limited. Chemosensory cues may rather allow conspecific social and partner recognition. We recorded the number of tongue-flick (TF) rates of T. wiegmanni amphisbaenians to scents of both sexes with different pairing social bonds (familiar vs. unfamiliar) presented on cotton swabs to test discrimination of social groups. As seen from a rise in the number of TFs, males discriminated unfamiliar females from unfamiliar males. This suggests that chemical cues may be used by males to locate new mates. In contrast, females detected scent of unfamiliar conspecifics, but did not show sex discrimination. Both males and females discriminated the scent of an individual with which they had formed a pair bond from an unfamiliar individual of the same sex as the partner. Also, males, but not females, were capable of self-recognition, suggesting that scent marks of males in home ranges may provide individual information in intrasexual relationships. We conclude that conspecific discrimination based on chemical cues may allow the maintenance of social relationships and relatively stable pairs in fossorial reptiles inhabiting visually restricted environments.

Increased temperature disrupts chemical communication in some species but not others: The importance of local adaptation and distribution.

Environmental conditions experienced by a species during its evolutionary history may shape the signals it uses for communication. Consequently, rapid environmental changes may lead to less effective signals, which interfere with communication between individuals, altering life history traits such as predator detection and mate searching. Increased temperature can reduce the efficacy of scent marks released by male lizards, but the extent to which this negative effect is related to specific biological traits and evolutionary histories across species and populations have not been explored. We experimentally tested how increased temperature affects the efficacy of chemical signals of high‐ and low‐altitude populations of three lizard species that differ in their ecological requirements and altitudinal distributions. We tested the behavioral chemosensory responses of males from each species and population to male scent marks that had been incubated at one of two temperatures (cold 16°C or hot 20°C). In high‐altitude populations of a mountain species (Iberolacerta monticola), the efficacy of chemical signals (i.e., latency time and number of tongue flicks) was lower after scent marks had been exposed to a hot temperature. The temperature that scent marks were incubated at did not affect the efficacy of chemical signals in a ubiquitous species (Podarcis muralis) or another mountain species (I. bonalli). Our results suggest that specific ecological traits arising through local adaptation to restricted distributions may be important in determining species vulnerability to climatic change.

Laboratory Demonstrations of Pheromone-Mediated Scent-Marking, Orientation, and Mounting Behavior in Polistes exclamans (Hymenoptera: Vespidae)

A combination of arena,Y-tube olfactometer, and flight tunnel assays were used to determine responses of male and reproductive female Polistes exclamans Vierick (Hymenoptera: Vespidae) to odors and extracts of conspecific males and females, as potential pheromone-mediated sexual behaviors. Males rubbed the sternites of the gaster on filter paper in response to extracts of the female mesosoma and gaster, indicating possible scent-marking. In a Y-tube olfactometer, females oriented to extracts of the male head, gaster, legs, seventh sternite, and mandibular glands. In the olfactometer, males oriented to extracts of the female whole body, but were repelled by extracts of the female head. Both females and males oriented to solvent washes of glass jars that held wasps of the opposite sex, as well as volatile collections from the opposite sex. In a flight tunnel, males and females both exhibited chemoanemotaxis that consisted of upwind flight and close range casting in response to odor piped into the tunnel from live wasps of the opposite sex. Males mounted a female wasp model treated with an extract of the female mesosoma. Together, these experiments suggest pheromones stimulate male scent-marking, attraction of females to males and males to females, and mounting, in P. exclamans. Results indicate the mesosoma as a source of female pheromones, and the legs, gastral sternal glands, and mandibular glands, as male pheromone sources.

Interpopulational Variations in Sexual Chemical Signals of Iberian Wall Lizards May Allow Maximizing Signal Efficiency under Different Climatic Conditions.

Sexual signals used in intraspecific communication are expected to evolve to maximize efficacy under a given climatic condition. Thus, chemical secretions of lizards might evolve in the evolutionary time to ensure that signals are perfectly tuned to local humidity and temperature conditions affecting their volatility and therefore their persistence and transmission through the environment. We tested experimentally whether interpopulational altitudinal differences in chemical composition of femoral gland secretions of male Iberian wall lizards (Podarcis hispanicus) have evolved to maximize efficacy of chemical signals in different environmental conditions. Chemical analyses first showed that the characteristics of chemical signals of male lizards differed between two populations inhabiting environments with different climatic conditions in spite of the fact that these two populations are closely related genetically. We also examined experimentally whether the temporal attenuation of the chemical stimuli depended on simulated climatic conditions. Thus, we used tongue-flick essays to test whether female lizards were able to detect male scent marks maintained under different conditions of temperature and humidity by chemosensory cues alone. Chemosensory tests showed that chemical signals of males had a lower efficacy (i.e. detectability and persistence) when temperature and dryness increase, but that these effects were more detrimental for signals of the highest elevation population, which occupies naturally colder and more humid environments. We suggest that the abiotic environment may cause a selective pressure on the form and expression of sexual chemical signals. Therefore, interpopulational differences in chemical profiles of femoral secretions of male P. hispanicus lizards may reflect adaptation to maximize the efficacy of the chemical signal in different climates.

The scent of sun worship: basking experience alters scent mark composition in male lizards

Signals used in female choice should honestly advertise the benefits that males can provide, with direct benefits often argued as being more important than indirect benefits. However, the nature of direct benefits in species without paternal care or nuptial gifts is poorly understood. Previous studies on lizards suggest that females decide where to settle and assumedly who to mate with based on information contained in scent marks from territorial males. Access to high-quality thermal resources is crucial for female reproductive success. Females may therefore be able to detect and exploit thermal-induced variation in the chemical composition of male scent marks when assessing the quality of his territory. We show that the amount of time male wall lizards (Podarcis muralis) are allowed to bask significantly alters the chemical composition of their femoral secretions used in scent marking. The direction of the change is consistent with adaptive plasticity to maintain signalling efficacy under warm conditions that increase evaporation of femoral secretions. The compounds affected by basking experience included those previously associated with male quality or shown to mediate male–male competition in lizards. However, whilst female lizards could discriminate between scent marks of males that had experienced different basking conditions, they did not preferentially associate with the scent from males from high-quality thermal conditions. These results highlight the potential importance of a previously neglected environmental effect on chemical signalling. We suggest thermal effects may have significant consequences for scent-mark composition in variable environments, with potential repercussions on olfactory communication in lizards.

Reintroducing domesticated wild mice to sociality induces adaptive transgenerational effects on MUP expression

When brought into captivity, wild animals can adapt to domestication within 10 generations. Such adaptations may decrease fitness in natural conditions. Many selective pressures are disrupted in captivity, including social behavioral networks. Although lack of sociality in captivity appears to mediate domestication, the underlying mechanisms are not well understood. Additionally, determining the contribution of genetic inheritance vs. transgenerational effects during relaxed selection may provide insight into the flexibility of adaptation. When wild-derived mice kept under laboratory conditions for eight generations were reintroduced to sociality and promiscuity (free mate choice), they adapted within two generations. Fitness assessments between this promiscuous lineage and a monogamous laboratory lineage revealed male-specific effects. Promiscuous-line males had deficits in viability, but a striking advantage in attracting mates, and their scent marks were also more attractive to females. Here, we investigate mechanistic details underlying this olfactory signal and identify a role of major urinary protein (MUP) pheromones. Promiscuous-line males inherit higher MUP expression than monogamous-line males through transgenerational inheritance. Sociality-driven maternal and paternal effects reveal intriguing conflicts among parents and offspring over pheromone expression. MUP up-regulation is not driven by hormone-driven transduction pathways, but rather is associated with reduction in DNA methylation of a CpG dinucleotide in the promoter. This reduction in methylation could enhance transcription by promoting the binding of transcription factor USF1 (upstream stimulatory factor 1). Finally, we experimentally demonstrate that increased MUP expression is a female attractant. These results identify molecular mechanisms guiding domestication and adaptive responses to fluctuating sociality.

Scent marking increases male reproductive success in wild house mice

Scent marking is often assumed to be a secondary sexual trait that increases males' mating and reproductive success, although direct evidence for this hypothesis is lacking. We conducted a study with wild-derived house mice, Mus musculus musculus, to test whether scent marking increases males' reproductive success when females can freely choose between two territorial males. We also experimentally manipulated males' competitive scent marking by exchanging scent-marked tiles between the neighbouring males' territories (intrusion treatment) or relocating males' tiles within their own territory (control). Experimental animals were tested twice and we examined whether individual males were consistent in their marking. We found that males marked more in the intrusion treatment than controls and more at shared territorial borders than elsewhere. We found high day-to-day variation in most individuals' scent marking, and yet the sum of individuals' scent marking was consistent over time and across different social conditions. Genetic paternity analyses revealed that males' scent marking significantly increased their reproductive success in both the intrusion treatment and the controls. Surprisingly, however, female social preference was not positively correlated with male scent marking. These results provide direct evidence that scent marking increases males' reproductive success when females can choose their mates, even though it did not increase females' social preferences.

Female mate choice based on pheromone content may inhibit reproductive isolation between distinct populations of Iberian wall lizards

Abstract The Iberian wall lizard Podarcis hispanica forms part of a species complex with several morphologically and genetically distinct types and populations, which may or may not be reproductively isolated. We analyzed whether female mate choice based on males’ chemical signals may contribute to a current pre-mating reproductive isolation between two distinct populations of P. hispanica from central Spain. We experimentally examined whether females choose to establish territories on areas scent-marked by males of their own population, versus areas marked by males of the other population. Results showed that females did not prefer scent-marks of males from their own population. In contrast, females seemed to attend mostly to among-individual variation in males’ pheromones that did not differ between populations. Finally, to test for strong premating reproductive isolation, we staged intersexual encounters between males and females. The population of origin of males and females did not affect the probability nor the duration of copulations. We suggest that the different environmental conditions in each population might be selecting for different morphologies and different chemical signals of males that maximize efficiency of communication in each environment. However, females in both populations based mate choice on a similar condition-dependent signal of males. Thus, male signals and female mate choice criteria could be precluding premating reproductive isolation between these phenotypically “distinct” populations.

Responses of female rock lizards to multiple scent marks of males: Effects of male age, male density and scent over-marking

Scent-marked substrates may inform conspecifics on the characteristics of territorial males. Scent-marks of male Carpetan rock lizards (Iberolacerta cyreni) affect space use of females, which by selecting an area may increase the probability of mating with the male that has scent-marked that area. However, males do not hold exclusive territories, and scent-marks of different individual males are often together. This may provide complex information from multiple sources on the social structure. Here, we examined female preference in response to scent marks of various males and combinations in a laboratory experiment. Females preferred areas scent-marked by territorial old males against those scent-marked by young satellite-sneaker males. This reflected the known preference of females for mating with old males. In a second experiment, females preferred areas scent-marked by two males to areas of similar size marked by a single male. This may increase the probability of obtaining multiple copulations with different males, which may favour sperm competition and cryptic female choice, or may be a way to avoid infertile males. Finally, when we experimentally over-marked the scent-marks of an old male with scent-marks of a young male, females did not avoid, nor prefer, the over-marked area, suggesting that the quality of the old male may override the presence of a satellite male. We suggest that, irrespective of the causes underlying why a female selects a scent-marked area, this strategy may affect her reproductive success, which may have the same evolutionary consequences that “direct” mate choice decisions of other animals.

Social behavior, chemical communication, and adult neurogenesis: Studies of scent mark function in Podarcis wall lizards

Lacertid lizards have been hailed as a model system for the study of reptilian chemical communication. However, results obtained with the genus Podarcis, a diverse group of wall lizards with complex systematics, challenge emerging paradigms and caution against hasty generalizations. Here we review the available evidence on the role of chemical stimuli in male–female and male–male interactions in Iberian Podarcis. Males of several species can discriminate between chemicals left on substrates by females of their own or a different species, suggesting that differences in female chemical cues may underlie species recognition in this group. Females, on the other hand, do not respond differentially to conspecific and congeneric male scent marks. Males of Podarcis liolepis use scent marks to recognize rivals individually, evaluate their competitive ability (i.e., body size), and assess the threat posed by each individual rival neighbor. In contrast, females do not exhibit a preference for territories scent marked by larger (i.e., more competitive) males, which suggests a limited role for male scent marks in pre-copulatory mate choice. This behavioral sex difference is consistent with detailed neuro-ethological evidence showing that chemosensory brain areas in P. liolepis are sexually dimorphic. The accessory olfactory bulbs are larger (both in absolute and relative terms) in males than in females, probably as a result of sex-specific rates of adult neurogenesis. In both sexes, cell proliferation undergoes seasonal cycles that may have evolved to satisfy increased chemosensory demands at particular times of the year. Overall, and against recent generalizations, these results suggest that male scent marks have been shaped mainly by strong intrasexual selection.

A Comparison of Scent Marking between a Monogamous and Promiscuous Species of Peromyscus: Pair Bonded Males Do Not Advertise to Novel Females

Scent marking can provide behavioral and physiological information including territory ownership and mate advertisement. It is unknown how mating status and pair cohabitation influence marking by males from different social systems. We compared the highly territorial and monogamous California mouse (Peromyscus californicus) to the less territorial and promiscuous white-footed mouse (P. leucopus). Single and mated males of both species were assigned to one of the following arenas lined with filter paper: control (unscented arena), male scented (previously scent-marked by a male conspecific), or females present (containing females in small cages). As expected, the territorial P. californicus scent marked and overmarked an unfamiliar male conspecific's scent marks more frequently than P. leucopus. Species differences in responses to novel females were also found based on mating status. The presence of unfamiliar females failed to induce changes in scent marking in pair bonded P. californicus even though virgin males increased marking behavior. Pair bonding appears to reduce male advertisement for novel females. This is in contrast to P. leucopus males that continue to advertise regardless of mating status. Our data suggest that communication through scent-marking can diverge significantly between species based on mating system and that there are physiological mechanisms that can inhibit responsiveness of males to female cues.

SOCIAL BEHAVIOR, CHEMICAL COMMUNICATION, AND ADULT NEUROGENESIS: STUDIES OF SCENT MARK FUNCTION IN IBERIAN PODARCIS WALL LIZARDS

Event Abstract Back to Event SOCIAL BEHAVIOR, CHEMICAL COMMUNICATION, AND ADULT NEUROGENESIS: STUDIES OF SCENT MARK FUNCTION IN IBERIAN PODARCIS WALL LIZARDS Enrique Font1* 1 Instituto Cavanilles, Universidad de Valencia, Spain Iberian lacertid lizards have been hailed as a model system for the study of reptilian chemical communication. However, results obtained with the genus Podarcis, a diverse group of wall lizards with complex systematics, challenge emerging paradigms and caution against hasty generalizations. Males of several species can discriminate between chemicals left on substrates by females of their own or a different species, suggesting that differences in female chemical cues may underlie species recognition in this group. Females, on the other hand, do not respond differentially to conspecific and congeneric male scent marks. This sex difference agrees with the finding that chemosensory brain areas in Podarcis are sexually dimorphic. The accessory olfactory bulbs are larger (both in absolute and relative terms) in males than in females, probably as a result of sex-specific rates of adult neurogenesis: males produce more new cells in telencephalic chemosensory brain areas (olfactory bulbs, lateral cortex, nucleus sphericus) than females. In both sexes, cell proliferation peaks in November-December (non-breeding season) and in April-May (breeding season). These seasonal cycles of adult neurogenesis may have evolved to satisfy increased chemosensory demands at particular times of the year. Male scent marks mediate important aspects of male-male competition and intraspecific territorial behavior. In P. liolepis (formerly P. hispanica), males use scent marks to recognize rivals individually, evaluate their competitive ability (i.e. body size), assess the threat posed by each individual rival neighbor, and probably even extract functional information about the quality of a rival’s territory. In contrast, females do not exhibit a preference for territories scent marked by larger (i.e. more competitive) males. These results suggest that male scent marks play a limited role in precopulatory mate choice and have been shaped instead by strong intrasexual selection. Keywords: chemical communication, Lizard, mate choice, mating system, pheromone, species recognition, visual communication Conference: ISAREN 2011: 7th International Symposium on Amphibian and Reptilian Endocrinology and Neurobiology, Ann Arbor, United States, 11 Jul - 13 Jul, 2011. Presentation Type: Invited Symposium Topic: Brain and behavior Citation: Font E (2011). SOCIAL BEHAVIOR, CHEMICAL COMMUNICATION, AND ADULT NEUROGENESIS: STUDIES OF SCENT MARK FUNCTION IN IBERIAN PODARCIS WALL LIZARDS. Front. Endocrinol. Conference Abstract: ISAREN 2011: 7th International Symposium on Amphibian and Reptilian Endocrinology and Neurobiology. doi: 10.3389/conf.fendo.2011.03.00011 Copyright: The abstracts in this collection have not been subject to any Frontiers peer review or checks, and are not endorsed by Frontiers. They are made available through the Frontiers publishing platform as a service to conference organizers and presenters. The copyright in the individual abstracts is owned by the author of each abstract or his/her employer unless otherwise stated. Each abstract, as well as the collection of abstracts, are published under a Creative Commons CC-BY 4.0 (attribution) licence (https://creativecommons.org/licenses/by/4.0/) and may thus be reproduced, translated, adapted and be the subject of derivative works provided the authors and Frontiers are attributed. For Frontiers’ terms and conditions please see https://www.frontiersin.org/legal/terms-and-conditions. Received: 26 Jul 2011; Published Online: 09 Aug 2011. * Correspondence: Dr. Enrique Font, Instituto Cavanilles, Universidad de Valencia, Paterna, Valencia, 46100, Spain, enrique.font@uv.es Login Required This action requires you to be registered with Frontiers and logged in. To register or login click here. Abstract Info Abstract The Authors in Frontiers Enrique Font Google Enrique Font Google Scholar Enrique Font PubMed Enrique Font Related Article in Frontiers Google Scholar PubMed Abstract Close Back to top Javascript is disabled. Please enable Javascript in your browser settings in order to see all the content on this page.

Responses to intruder scents in the cooperatively breeding meerkat: sex and social status differences and temporal variation

Although sex-specific responses to intruder scent marks have been described in many mammal species, less is known about variation in responses in cooperatively breeding species where differential access to breeding opportunities exists within groups. When reproductive skew is high, strong responses to extragroup breeding rivals would be expected mainly from dominant individuals, with levels of investment depending on female receptiveness. However, evidence from controlled experiments on cooperative breeders for sex and social status differences in responses to intruder scent marks is limited. Here, we investigated responses to intruder scent marks in cooperatively breeding meerkats (Suricata suricatta), where a single dominant pair largely monopolizes within-group reproduction. A series of experimental presentations using feces were used to test first, whether meerkats discriminate between resident and extragroup male scent marks; second, whether sex and social status affect the response to intruding male scents; and third, whether dominant males increase their level of response when dominant females are most receptive. Our results suggest that meerkats are able to discriminate between resident and intruding male scent marks and show that dominant males have the strongest overall response to intruder scent marks, which does not increase with female receptiveness. We suggest that, although all group members may be affected by the presence of intruders, reproductive conflict may be the main reason for the stronger response of dominant males to extragroup male scent marks in this cooperatively

The role of scent marks in female choice of territories and refuges in a lizard (Podarcis hispanica).

Female mate choice based on male phenotypic traits is controversial in lizards, particularly in territorial species. In this study, we examine female choice of male scent marks in a territorial lacertid lizard (Podarcis hispanica) in which scent marks have been shown to signal male size (i.e., an important determinant of competitive ability in this species). Females were simultaneously exposed to three naturalized 4 m² choice areas bearing: (1) no scent marks (i.e., control), (2) scent marks of large males, and (3) scent marks of small males. Although female lizards preferentially associated with scent marked choice areas, we found no evidence that females chose territories marked by large males. Furthermore, in response to experimentally induced dusk at the end of choice trials, females preferentially took shelter in refuges scent marked by small males. Our results suggest that, like males in this species, females are able to use male scent marks to assess the body size of resident territorial males, but do not show a preference for territories occupied by large males.

Female urine-induced male mice ultrasonic vocalizations, but not scent-marking, is modulated by social experience

Despite the evidence for a communicative function of rodent scent marks and ultrasonic vocalizations, relatively little is known about the impact of social factors on these two forms of communication. Here, we tested the effects of two important social factors, prior exposure to a female and freshness of female urine, on male scent marks and ultrasonic vocalizations elicited by female urine. We also asked whether a recently reported strain difference between the highly social strain C57BL/6J (B6) and the mouse model of autism BTBR T+tf/J (BTBR) herein is specifically seen in response to female urine or also detectable in response to male urine traces. Results show that the emission of female urine-elicited ultrasonic vocalizations was dependent on previous female experience, while scent-marking behavior was not affected. A positive correlation was detected between scent-marking behavior and ultrasonic calling in the most biologically relevant context, male mice exposed to fresh female urine after female experience. Correlations were less prominent or missing in less biologically relevant contexts, e.g. in male mice exposed to fresh female urine without previous female experience, indicating that previous female experience is affecting both the emission of female urine-elicited ultrasonic vocalizations and the correlation between olfactory and acoustic communication. The strain difference in scent-marking behavior and ultrasonic calling between B6 and BTBR appears to be specific to female urine-elicited behavior as it was not seen in response to male urine traces, highlighting the relevance of the social context in which mouse communication is evaluated.

Temporal and Spatial Distribution of Male Scent Marks in the Polygynous Greater Sac-Winged Bat

Scent marks are relatively long-lived signals that can be perceived by conspecifics when the producer is absent. Therefore, it is often not obvious to whom the signal is directed. In daytime roosts of the polygynous greater sac-winged bat, males scent mark territories with facial gland secretions. Territories are a valuable resource for males, as they offer exclusive courtship opportunities, which results in increased male reproductive success and, consequently, increased male-male competition over territories. The information encoded in male scent marks could, therefore, be either directed at females as part of an olfactory courtship display or at male competitors as part of territorial behaviour. We expected territorial males to scent mark in the morning, shortly before females return to the territory and close to female roosting sites, if scent marks are directed at females as part of the courtship display. And we expected harem males to scent mark at the territory boundaries, where male-male encounters are most likely to occur, if scent marks are directed at male competitors. We found that males marked more frequently in the afternoon, at a time when all females have already left the territory, and harem males marked at the territory boundaries and not inside their territory in the area where females roost. At boundaries males fan volatiles from specialised wing sacs towards competitors outside the territory. Scent marking of male Saccopteryx bilineata might therefore be congruent with the assessment-hypothesis, which states that scent marks offer intruders the possibility to make an olfactory assessment of the territory owner without direct physical interaction. Thus, scent marks of male S. bilineata are most likely influenced by male-male competition and not by female choice.

Behavioral and reproductive responses of female opossums to volatile and nonvolatile components of male suprasternal gland secretion

Female gray short-tailed opossums ( Monodelphis domestica) lack an estrous cycle and are induced into estrus by exposure to a pheromone in male scent marks. Behavioral and physiological responses of females to the volatile and nonvolatile components of scent marks were examined in two experiments. Young females ( n = 9) were tested prior to and during their first estrus for behavioral responses to scent marks, collected on a 7-ml glass vial rubbed over the suprasternal gland of a mature male. The response to volatile components of the scent mark, recorded when marked and unmarked vials were covered with a perforated shield, was compared to the response to these vials when unshielded. Estrous females nuzzled the shields over marked vials (55.8 ± 8.5 nuzzles/10 min) more than the shielded clean vial (10.9 ± 2.4) ( P < 0.05); a similar response was observed in anestrous females. Nuzzling of unshielded, scent-marked vials was higher ( P < 0.05) during anestrus than in the same females when in estrus. The role of nonvolatile pheromones in reproductive activation was tested in adult females ( n = 11) exposed for up to 14 days to a shielded, marked vial or to an unshielded, marked vial in a crossover design. All females exposed to unshielded vials expressed estrus, and 10 copulated. Only 2 females expressed estrus (significantly fewer, P < 0.05), when exposed to shielded marked vials, and neither copulated. These results demonstrate that females detect and respond behaviorally to both volatile and nonvolatile components of male suprasternal gland secretion, but the estrus-inducing pheromone in these secretions is nonvolatile.

Female sensory bias may allow honest chemical signaling by male Iberian rock lizards

Some mate choice theories propose that only male signals that are honest and condition-dependent can be stable, while another hypothesis states that males evolve signals that exploit the sensory system of females. However, sensory traps might evolve into honest signals if they are differentially costly for males. We tested whether a pre-existing sensory bias for food chemicals explained chemosensory preferences of female Iberian rock lizards for male scents. We manipulated hunger levels of females and found that food-deprived females had increased chemosensory responses to chemical stimuli from both invertebrate prey and femoral secretions of males, but not to control water. Further tests suggested that cholesta-5,7-dien-3-ol (provitamin D3), a lipid found in both prey and males’ scent, may be one of the chemicals eliciting these responses. Moreover, hungry females spent more time on scent marks of males that had experimentally increased cholesta-5,7-dien-3-ol than on scent marks of males alone, whereas for control females this effect was not significant. We suggest that preexisting sensory bias for essential nutrients (i.e., provitamin D) may be the origin of similar female responses to male chemicals. However, previous studies have suggested that the allocation of these chemicals to ornaments is costly and only high quality males can afford it. Therefore, preexisting sensory bias for essential nutrients may further allow the evolution and maintenance of honest sexual displays.