Olfactory Experience Affects the Response of Meadow Voles to the Opposite‐Sex Scent Donor of Mixed‐Sex Over‐Marks

Scent marking and over‐marking are important forms of communication between the sexes for many terrestrial mammals. Over the course of three experiments, we determined whether the amount of time individuals investigate the scent marks of opposite‐sex conspecifics is affected by 4 d of olfactory experience with those conspecifics. In Experiment 1, female meadow voles, Microtus pennsylvanicus, spent more time investigating the scent mark of the novel male conspecific than that of the familiar male donor, whereas male voles spent similar amounts of time investigating the scent mark of the familiar female and a novel female conspecific. In Experiment 2, voles were exposed to a mixed‐sex over‐mark in which subjects did not have 4 d of olfactory experience with either the top‐scent donor or the bottom‐scent donor. During the test phase, male and female voles spent more time investigating the scent mark of the opposite‐sex conspecific that provided the top‐scent mark than that of a novel, opposite‐sex conspecific. Male and female voles spent similar amounts of time investigating the scent mark of the bottom‐scent donor and that of a novel opposite‐sex conspecific. In Experiment 3, voles were exposed to a mixed‐sex over‐mark that contained the scent mark of an opposite‐sex conspecific with which they had 4 d of olfactory experience. During the test phase, male voles spent more time investigating the mark of the familiar, top‐scent female than the scent mark of a novel female donor but spent similar amounts of time investigating the mark of the familiar, bottom‐scent female and that of a novel female donor. In contrast, female voles spent more time investigating the mark of a novel male donor than that of either the familiar, top‐scent male or that of the familiar, bottom‐scent male. The sex differences in the responses of voles to scent marks and mixed‐sex over‐marks are discussed in relation to the natural history and non‐monogamous mating system of meadow voles.     

Recognition of individual,sex and species odours by salamanders of the Plethodon glutinosus-P. Jordani complex

Salamanders of the Plethodon glutinosus-P. jordani complex were tested for the ability to distinguish conspecific, sex-specific, and heterospecific chemical cues. Male and female P. glutinosus preferred substrates previously occupied by conspecifics over their own, but randomly chose between substrates marked by male or female conspecifics. This suggests that while these salamanders are able to distinguish between their own and conspecific substrate odours, they are unable to identify sex by means of substrate odours. Experiments using an olfactometer showed that male P. jordani, male and female P. glutinosus, and an electrophoretically distinct and non-hybridizing sympatric phenotype in the P. glutinosus complex (here called species A), all preferred female airborne odours over male airborne odours. This demonstrates that these salamanders can identify sex by means of airborne odours. Male P. glutinosus and species A both preferred conspecific female odours over heterospecific female odours in olfactometer experiments. These results suggest an important role for olfaction in the sexual and social behaviour of these salamanders, particularly as a pre-mating isolating mechanism.     

Sex Discrimination via Anal Gland Secretion in a Territorial Monogamous Mammal

Monogamy in mammals is relatively unusual, only occurring in 3–5% and to an even lesser degree in rodents. Monogamy exists in two forms, facultative which evolved due to female dispersion and obligate when male care is needed for offspring survival. Most monogamous rodents use scent marking as a form of territorial defence to obtain exclusive access to vital resources. Intrasexual competition occurs in many species to maintain pair bonds and also to signal presence within a territory. In this study, we tested the hypothesis that resident obligate monogamous Eurasian beaver (Castor fiber) can discriminate between the sexes when investigating anal gland secretions (AGS) from unknown (stranger) mated pairs. We predicted that, due to intrasexual competition, dominant territorial male and female beavers will display a stronger response to AGS of same‐sex conspecifics. Territorial intrusion by mated pair strangers was simulated by the formation of experimental scent mounds (ESM) with AGS. Our results showed that both sexes (1) displayed a stronger response to the male AGS in sniffing duration, (2) physically responded for longer durations towards male AGS and (3) overmarked male AGS more frequently. We infer that obligate monogamous mammals can determine the sexual identity of intruding conspecifics of a mated pair via AGS and that intruding males may pose more of a threat to resident males via intrasexual competition and resident females due to the long‐term costs, such as infanticide and abortion, of a new dominant male.     

Effects of sex hormones and gender on attraction thresholds for volatile anal scent gland odors in ferrets

Body odors contribute to mate recognition and sexual partner preference in many mammals, including ferrets. We used a habituation/dishabituation procedure to test whether sex steroid hormones influence whether ferrets will approach and investigate different concentrations of volatile anal scent gland odors from male and female conspecifics. When tested with high concentrations of anal scent gland secretions in oil vehicle, gonadectomized male and female ferrets that received no sex steroids reliably discriminated anal scents from male and female conspecifics. This discrimination most likely reflects gender recognition rather than individual recognition because gonadectomized, sex steroid-treated ferrets discriminated between anal scents of males and females but not between anal scents of individual males or females. Treatment with either the estrogen receptor agonist, estradiol benzoate (EB), or the androgen receptor agonist, 5-alpha dihydrotestosterone proprionate (DHTP), increased investigation of low concentrations of anal scent by gonadectomized ferrets. These data suggest that ferrets could use anal scent gland secretions in mate recognition and that seasonal increases in circulating sex steroid hormones increase ferrets' responsiveness to low concentrations of these odors.     

Effects of vomeronasal organ removal on olfactory sex discrimination and odor preferences of female ferrets

Previous research suggests that body odorants, including anal scents and urinary odors, contribute to sex discrimination and mate identification in European ferrets of both sexes. We assessed the possible role of the vomeronasal organ (VNO) in these functions by surgically removing the organ bilaterally in sexually experienced female ferrets. Lesioned (VNOx) and sham-operated control (VNOi) females reliably discriminated between male- and female-derived anal scent gland as well as fresh urinary odors in habituation/dishabituation tests. However, VNOi females spent significantly more time than VNOx subjects investigating male urinary odors in these tests. Also, VNOi females, but not VNOx subjects, preferred to investigate day-old male versus female urine spots as well as wooden blocks that had previously been soiled by male versus female ferrets. Both groups of female ferrets preferred to approach volatile odors from a breeding male instead of an estrous female in Y-maze tests and both groups showed similar levels of receptive sexual behavior in response to a male's neck grip. The VNO is apparently not required for olfactory sex discrimination or mate recognition in this carnivore, but instead may play a role in promoting continued contact with nonvolatile body odors previously deposited by opposite-sex conspecifics during territorial scent marking.     

In Space and Time: Territorial Animals are Attracted to Conspecific Chemical Cues

Territorial animals lay scent marks around their territories to broadcast their presence, but these olfactory signals can both attract and repel conspecifics. Attraction or aversion can have a profound impact in terms of space use and thereby influence an individual's access to resources and mates. Here, we test the impact of chemical signals on the long‐term space use and activity of receivers, comparing the response of males and females, territory holders, and temporary visitors in Sceloporus undulatus lizards in the field. We placed either male femoral gland secretions (chemical) or blank (control) cues on resident male landmarks, repeatedly over 5 d, while monitoring the activity and location of all lizards in the vicinity. We found that resident males and females, but not non‐resident males, were active on more days near landmarks treated with chemical cues than landmarks treated with control cues. Non‐resident males remained closer to chemical than control cues. These results suggest that territorial scent marks are attractive to conspecifics and impact space use, but that the specific effects depend on receiver sex and residency status. Such subtle or gradual changes in behavior may frequently be overlooked by short‐term choice experiments. Future studies investigating the behavioral significance of a communicative signal should consider these finer details of behavior for a more comprehensive assessment.     

Evolutionary patterns of major urinary protein scent signals in house mice and relatives

Scent marks are important mediators of territorial behaviour and sexual selection, especially among mammals. The evolution of compounds used in scent marks has the potential to inform our understanding of signal evolution in relation to social and sexual selection. A major challenge in studies of chemical communication is that the link between semiochemical compounds and genetic changes is often unclear. The major urinary proteins (MUPs) of house mice provide information on sex, status and individual identity. Importantly, MUPs are a direct protein product of genes, providing a clear link between genotype and phenotype. Here, we examine the evolution of urinary protein signals among house mice and relatives by examining the sequences and patterns of mRNA expression of Mup genes related to urinary scent marks. MUP patterns have evolved among mouse species both by gene duplication and variation in expression. Notably, protein scent signals that are male specific in well‐studied inbred laboratory strains vary in sex‐specificity among species. Our data reveal that individual identity signals in MUPs evolved prior to 0.35 million years ago and have rapidly diversified through recombining a modest number of amino acid variants. Amino acid variants are much more common on the exterior of the protein where they could interact with vomeronasal receptors, suggesting that chemosensory perception may have played a major role in shaping MUP diversity. These data highlight diverse processes and pressures shaping scent signals, and suggest new avenues for using wild mice to probe the evolution of signals and signal processing.     

Male and Female Meadow Voles,Microtus pennsylvanicus,Differ in Their Responses to Heterospecific/Conspecific Over‐Marks

Voles use runways, paths, and trails that may also be used by rabbits and mink. These shared areas could contain the scent marks of conspecifics and heterospecifics. Thus, it is likely that the scent marks of heterospecifics may overlap or be overlapped by those of voles, forming over‐marks. Much is known about how voles respond to over‐marks of two different conspecifics. However, we do not know how they would respond to an opposite‐sex conspecific whose scent marks are in an over‐mark with the scent marks of predator or the scent marks of a non‐predator heterospecifics. We tested the hypothesis that meadow voles, Microtus pennsylvanicus, differ in their response to the scent mark of the opposite‐sex conspecific if the scent mark was overlapped by that of a mink, a vole predator, or rabbit, a vole non‐predator. We found that female but not male voles showed a preference for the scent marks of the opposite‐sex conspecifics that were part of the mink‐vole over‐mark when compared to those of opposite‐sex conspecifics that were not part of the over‐mark. This preference by female voles was independent of whether the male vole was the top‐scent donor or bottom‐scent donor of the over‐mark. Male and female voles showed no preference between the scent marks of the opposite‐sex conspecifics whose marks were part of or not part of the rabbit‐vole over‐mark. Sex differences in the manner that meadow voles respond to rabbit‐vole and mink‐vole over‐marks are discussed.     

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.     

The Reproductive State of Female Voles Affects their Scent Marking Behavior and the Responses of Male Conspecifics to Such Marks

During the breeding season, the reproductive condition of female mammals changes. Females may or may not be sexually receptive. We conducted a series of experiments to determine whether reproductive condition of female meadow voles affects their scent marking behavior as well as the scent marking behavior of male conspecifics. In expt 1, females in postpartum estrus (PPE females) deposited more scent marks than females that were neither pregnant nor lactating (REF females) or ovariectomized females (OVX females). In expt 2, male voles scent marked more and deposited more over‐marks in areas marked by PPE females than by REF and OVX females. In expt 3, PPE females deposited more scent marks and over‐marks in areas marked by males than did females in the other reproductive states. The results of these experiments showed that male and female voles may vary in the number, type and location of scent marks they deposit in areas scented by particular conspecifics.     

The Role of Wing Coloration in Sex Recognition and Competitor Recognition in Rubyspot Damselflies (Hetaerina spp.)

The decision rules that animals use for distinguishing between conspecifics of different age and sex classes are relevant for understanding how closely related species interact in sympatry. In rubyspot damselflies (Hetaerina spp.), the red wing coloration of mature males is hypothesized to be a key trait for sex recognition and competitor recognition within species and the proximate trigger for interspecific male–male aggression. We tested this hypothesis by manipulating the wing coloration of tethered conspecific intruders and measuring the responses of territory holders of three species in the field. As predicted, covering the red spots of mature males with black ink nearly eliminated territorial responses, and in some cases, territorial holders clasped the blackened males as if they were females. Adding red spots to female wings triggered territorial responses and nearly eliminated sexual responses. Immature males with artificial red spots were attacked at the same rate as mature male intruders, and much more frequently than were immature male controls. The results varied somewhat by species. In H. titia, the only species of Hetaerina with substantial black wing pigmentation, the effects of blackening the red spots of intruders varied both geographically and seasonally. But even when blackening the red spots of male intruders did not reduce the aggressive response of H. titia territory holders, adding artificial red spots to female wings elicited aggressive responses and nearly eliminated sexual responses. The results of this study further strengthen the evidence that interspecific aggression in Hetaerina results from overlap in territorial signals and that the derived black wing pigmentation of H. titia reduces interspecific aggression.     

Effects of prepubertal ovariectomy on the development of scent glands, scent marking, and aggressive behaviors of female tamarin monkeys (Saguinus fuscicollis)

The effects of prepubertal ovariectomy on scent gland development, scent marking behavior, and on social interactions with strange adult conspecifics were studied in adult females cohabiting with intact males. Eight females were ovariectomized and eight underwent control surgery at 6 months of age. All were returned to their families after surgery and allowed to reach sexual maturity. At 18 months of age, each female was permanently paired with an adult, intact male. When the subjects were 21 months old, the social interactions of all pairs with strange adults were tested in a situation analogous to a territorial encounter. The scent marking activities of the subjects and their mates were studied during “territorial encounters,” in trial-free control situations, and in the presence of novel objects. Ovariectomy prior to puberty retarded the development of the scent gland but did not inhibit it completely. Social interactions with strange conspecifics were also affected. Ovariectomized females showed fewer threat displays than did control females, but there was no significant difference in the amount of injurious aggression both female groups directed at the strangers. Under all testing conditions, ovariectomized females tended to scent mark less frequently with the circumgenital-suprapubic gland than did controls. This difference was statistically significant only under some of the testing conditions. Intact females scent marked more frequently than their males but ovariectomized females did so only under trial-free testing conditions. The sternal scent gland was used very infrequently by all subjects and there was no difference in marking activity between males and females or between ovariectomized and control females.     

Food Deprivation Suppresses a Preference for the Top-Scent Mark of an Over-Mark in Meadow Voles (Microtus pennsylvanicus)

Food availability affects whether mammals communicate their interest in interacting with opposite‐sex conspecifics. This study examined the responses of voles to over‐marks, and factors that influence the formation and maintenance of a preference for the top‐scent in an over‐mark. Specifically, we investigated how food deprivation affected the amount of time male and female voles exposed to an over‐mark, later responded to the marks of the top‐ and bottom‐scent donors when subsequently presented with the two scents side by side. Males and females that were not food deprived and males that were food deprived 6 h before exposure to an over‐mark later maintained a preference for the donor of the top‐scent mark compared with the donor of the bottom‐scent mark of the over‐mark. Females that were food deprived for 6 h before or after exposure of the over‐mark and males food deprived 6 h after the exposure to the over‐mark showed no preference for the top‐scent mark donor. Re‐feeding females that were food deprived for 6 h before exposure to an over‐mark was sufficient to restore their preference for the mark of the top‐scent male over that of the bottom‐scent male. The different responses of food‐deprived male and female voles to over‐marks of opposite‐sex conspecifics may be associated with differences in their tactics for interacting with potential mates and the higher energetic costs of reproduction in female voles than in male voles.     

Floral scent is different between sexual phases within individuals in a synchronously dichogamous shrub (Canella winterana) but there is no distinct female or male scent profile across individuals

Floral scent is a highly complex and variable floral characteristic that is involved in pollinator attraction. One possible cause for variation in floral scent can be sexual identity of the flower. Here, we examine the floral scent bouquet of a synchronously dichogamous shrub, Canella winterana (L.) Gaertn.We used dynamic headspace extraction and gas chromatography – mass spectroscopy, followed by statistical analysis using non-metric multidimensional scaling, SIMPER, MANOVA, and PERMANOVA to identify and compare the scent profiles of Canella winterana in its female and male phase for multiple individuals as well as multiple inflorescences of a single tree over one cycle of its entire sexual phase transition. The scent profile of C. winterana is composed of 49 volatile organic compounds and dominated by five compounds. We found no evidence for distinct male or female scent profiles; however, there were significant differences in scent emission between different sexes within some individuals. Two compounds explained over half of the variation between sexual stages within individuals. Our exploration of a single tree's sexual phase transition, including neuter phase, found that five compounds dominated the female phase scent bouquet and that female phase was distinct from male and neuter phase. This study offers new insight into the role that variability in floral scent between sexual phases might play in variable pollinator behavioral responses. These results suggest partial support for two distinct hypotheses regarding the differences between the sex phases (1) honest signaling and (2) sexual mimicry.     

Meadow voles and prairie voles differ in the percentage of conspecific marks they over-mark

Many terrestrial mammals scent mark in areas containing the scent marks of conspecifics, and thus, may deposit their own scent marks on top of those that were deposited previously by conspecifics. This phenomenon, known as over-marking appears to play a role in same-sex competition or mate attraction. The present study determines whether meadow and prairie voles avoid over-marking the scent marks of conspecifics, target the scent marks of conspecifics and over-mark them, or randomly over-mark the scent marks of conspecifics. The data show that meadow and prairie voles adjust the number and location of scent marks that they deposit in areas marked previously by particular conspecifics. Male and female meadow and prairie voles target the scent marks of opposite-sex conspecifics and over-mark them. Female meadow and prairie voles also target the scent marks of female conspecifics. In contrast, male meadow and prairie voles over-mark the scent marks of male conspecifics in a random manner. By differentially over-marking the scent marks of conspecifics, voles may be able to communicate particular information to a variety of conspecifics.     

The effects of sex, reproductive condition and context on discrimination of conspecific odours by giant pandas

To elucidate some of the functions of chemical communication in giant pandas, Ailuropoda melanoleuca, we systematically exposed captive males, oestrous and nonoestrous females to each others' odours by providing access to one another's temporarily unoccupied enclosures. In comparison with control observations, pandas from all reproductive categories displayed a dramatic increase in several measures of chemosensory responsiveness, both when visiting another panda's enclosure and when encountering scent deposited in their home enclosure. Evidence for discrimination between odours encountered as a resident versus a visitor was minimal, but male pandas' response patterns were suggestive of a territorial function. We also found evidence for discrimination of sex and reproductive condition via chemical cues. Males showed a marked preference for female odours, investigating, licking, scent marking and vocalizing more in response to female than male odours. Males also vocalized more in response to oestrous than nonoestrous female odours. Nonoestrous females licked more and oestrous females vocalized more when encountering male than female odours. Our data on vocalizations suggest a potential role for odours in the activation of sexual motivation. We argue that the low reproductive success observed in captivity may be attributed in part to failure to provide sufficient opportunities for chemical communication, and encourage the judicious management of social odours to promote natural mating (e.g. using intersexual odour exposure to increase libido and decrease aggressiveness prior to mating introductions). Copyright 2000 The Association for the Study of Animal Behaviour.     

Social Reversal of Sex‐Biased Aggression and Dominance in a Biparental Cichlid Fish (Julidochromis marlieri)

In biparental species, aggression, dominance, and parental care are typically sexually dimorphic. While behavioral dimorphism is often strongly linked to gonadal sex, the environment—either social or ecological—may also influence sex‐biased behavior. In the biparental cichlid fish Julidochromis marlieri, the typical social environment for breeding pairs consists of large females paired with smaller males. While both sexes are capable of providing territory defense and parental care, the larger female provides the majority of defense for the pair, while the smaller male remains in the nest guarding their offspring. We examine the contributions of sex and relative mate size to these sex‐biased behaviors in monogamous J. marlieri pairs. Both female‐larger and male‐larger pairs were formed in the laboratory and were observed for territorial aggression (against conspecifics and heterospecifics), dominance, and parental care. In female‐larger pairs, territorial aggression and intra‐pair dominance were female‐biased, while in male‐larger pairs this bias was reversed. For both pairing types, the presence of an intruder amplified sex differences in territorial aggression, with the larger fish always attacking with greater frequency than its mate. Though less robust, there was evidence for plasticity of sex‐bias for some egg care related behaviors in the inverse direction. Our study suggests that relative mate size strongly influences the sex bias of aggression and dominance in J. marlieri and that this aspect of the social environment can override the influence of gonadal sex on an individual's behavior. The remarkable plasticity of this species makes Julidochromis an exciting model that could be used to address the relationship between proximate and ultimate mechanisms of behavioral plasticity.     

Lack of Evidence for the Prior Residence Effect in the Allegheny Mountain Dusky Salamander (Desmognathus ochrophaeus)

Territorial disputes are frequently settled by an advantage afforded to one of the contestants by asymmetries such as size difference, strength and motivation. Allegheny Mountain Dusky Salamanders (Desmognathus ochrophaeus) are reported to defend cover objects, a form of territorial behaviour. We conducted an experiment to determine whether or not adult salamanders of this species exhibit prior residence effect during staged encounters involving size‐matched, same sex conspecifics (i.e. does familiarity with a territory predict successful defence?). We tested 154 adult salamanders (72 female, 82 male) in reciprocal trials in which each animal acted as a resident and as an intruder. We recorded multiple agonistic behaviours including: front‐trunk raised, all‐trunk raised, nudge, bite, bite‐and‐hold and jaw‐lock. We assigned these behaviours scaled point values and calculated an index of aggression for each animal as a resident and as an intruder. We found that in same sex trials, males were significantly more aggressive than females. Although D. ochrophaeus exhibited stereotypical, agonistic behaviours similar to those reported for Plethodon, unlike Plethodon the outcomes of symmetrical social encounters were not influenced by residential status. Prior residence is documented to be a major determinant of territorial disputes among diverse animal taxa, including salamanders. However, our research suggests that the phenomenon is not universal and may depend on qualities of the microhabitat in which a species has evolved.     

Self‐Grooming Response of Meadow Voles to the Odor of Opposite‐Sex Conspecifics in Relation to the Dietary Protein Content of Both Sexes

Many animals self‐groom when they encounter the scent marks of opposite‐sex conspecifics. Self‐grooming transmits odiferous substances that contain information about the groomer’s condition, which is affected by its nutritional state. We tested the hypothesis that the amount of time that individuals self‐groom to opposite‐sex conspecifics is affected by the amount of protein in their diet and that of the scent donor. We did so by feeding meadow voles (Microtus pennsylvanicus) a diet containing 9%, 13%, or 22% dietary protein for 30 d and observing their self‐grooming behavior when they were exposed to bedding scented by an opposite‐sex conspecific (odor donor) fed one of the three diets, or fresh cotton bedding (control). The hypothesis was partially supported. We found that the protein content of the diet of male and female groomers did not affect the amount of time they self‐groomed. However, the protein content of the diet of male odor donors affected the amount of time that female voles spent self‐grooming. Female voles self‐groomed more in response to male odor donors fed a 22% protein‐content diet than to those produced by male odor donors fed either a 9% or a 13% protein‐content diet. Interestingly, the amount of time males self‐groomed was not affected by the protein content of the diet of the female odor donor. These results may, in part, be explained by the natural history of free‐living meadow voles, sex differences in costs associated with mate attraction and reproduction, and the direct or indirect benefits that females receive from males fed a diet high in protein content.     

Scent Marking in Female Prairie Voles: a Test of Alternative Hypotheses

We conducted three experiments with females in different stages of reproductive condition to test alternative hypotheses for the function of scent marking in female prairie voles, Microtus ochrogaster . The three reproductive categories were isolated females prior to sexual stimulation (anoestrous), sexually stimulated (oestrous) and lactating. Females in different reproductive condition were given the opportunity to scent mark clean unmarked substrate or areas that had previously been marked by adult females or adult males. The numbers of scent marks deposited by females did not differ statistically for females in different reproductive condition. However, there was a trend for anoestrous females to mark the most, oestrous females less, and lactating females the least. The lack of scent marking by lactating females might be to reduce conspicuousness to conspecifics or predators. Oestrous females tended to mark the most in the area marked previously by males, although the difference was not statistically significant.
Our results provide some support for a mate-attraction hypothesis and a territorial-defense hypothesis, but were most consistent with a self-advertisement hypothesis. Over marking was uncommon and did not differ by experiment or sex of previous donor. Our results suggest that the number and placement of scent marks by females are highly variable and function primarily to convey individual identity.