Scent-marking Rates Research Articles

Territorial scent-marking and proestrus in a recolonizing wild Gray Wolf (<i>Canis lupus</i>) population in central Wisconsin

Gray Wolf (Canis lupus) uses scent-marking to communicate breeding status, dominance, and territorial boundaries. Despite its importance for reproduction and pack dynamics, information on scent-marking and proestrus in wild wolf populations is limited to a handful of locations. We estimated the rate of territorial scent-marking and the probability of proestrus in a recolonizing Gray Wolf population near the species southern range extent in eastern North America. An analysis of 221 pack-winters of tracking data show that the incremental addition of one wolf pack increased marking rates by 3.4%, whereas increasing the number of wolves in a pack decreased marking rates by 12.1%. Scent-marking rates subsequently increased from 1.9 times/km during recolonization to 3.0 times/km once the population was saturated. We observed evidence of proestrus from 19 December to 14 March with the highest probability of proestrus occurring around 6 February, after peak marking rates around 26 January. Repeated observations of bloody urinations within individual packs suggest proestrus averages 27.9 days. Our study reveals the role of population growth on territorial behaviours and provides a foundation for studies exploring the role of geographic and temporal variation on territorial and reproductive behaviours in wolves.

Marking Versus Overmarking: Spatial and Behavioral Patterns of Scent Marking in Wild Diademed Sifaka (Propithecus diadema)

In mammals, olfactory communication plays an essential role in territorial and mating dynamics. Scent depositions in various species, including lemurs, can be placed via marking or overmarking (marking over previous depositions). We focused on the role that marking and overmarking play in territorial defence and intrasexual competition. We investigated these aspects in diademed sifaka (Propithecus diadema) in the primary rainforest of Maromizaha (eastern Madagascar). We collected scent marking data for five groups from April to November 2018 and from May to December 2019. We aimed to understand whether the lemurs deposited scent marks homogeneously across the home range and whether sex, rank, and occurrence of intergroup encounters affected the lemur’s deposition rate. We also asked whether males overmarked adult females more often than other depositions, and the marking and overmarking rates changed between the migration and non-migration seasons. We found that scent marking was performed higher in peripheral and overlapping areas than in the home range central areas. In addition, males had higher scent marking rates, but intergroup encounters did not affect deposition rates. Males showed higher rates of overmarking and primarily targeted dominant females’ depositions, particularly during the “migration” season (including premating and mating seasons). Our findings suggest a border-marking strategy in Propithecus diadema. More frequent scent marking in the “migration” season suggests intrasexual competition in males. Our results suggest that marking is associated with territorial and resource defence, suggesting that it plays a role in monopolizing females using a mate-guarding strategy and may also serve for males’ self-advertisement to females and subordinate depositors.

The potential interplay between the glandular microbiome and scent marking behavior in owl monkeys (Aotus nancymaae).

In mammals, scent marking behavior is a pervasive form of chemical communication that regulates social interactions within and between groups. Glandular microbiota consist of bacterial communities capable of producing chemical cues used in olfactory communication. Despite countless studies on scent marking in primates, few have examined the microbiota associated with glandular secretions. Nancy Ma's owl monkeys (Aotus nancymaae) are nocturnal, socially monogamous primates that frequently scent mark using their subcaudal glands. Previous analyses revealed that unique chemical signatures of Aotus may convey information about sex and age. We used positive reinforcement to sample the subcaudal glands of 23 captive owl monkeys to describe their glandular microbiomes and examine how patterns in these bacterial communities vary with age, sex, rearing environment and/or social group (pair identity). We coupled these analyses with behavioral observations to examine patterns in their scent marking behavior. We isolated 31 bacterial species from Phyla Firmicutes, Proteobacteria, and Actinobacteria, consistent with the dermal and glandular microbiomes of other primates. Several bacterial taxa we identified produce volatile organic compounds, which may contribute to olfactory communication. These bacterial communities are best predicted by an interaction between sex, rearing environment and pair identity rather than any of these variables alone. Within mated pairs of A. nancymaae, males and females scent mark their nest boxes at similar frequencies. In some pairs, rates of scent marking by males and females fluctuated over time in a similar manner. Pairs that had been together longer tended to exhibit the greatest similarities in their rates of scent marking. Together, these findings suggest that scent marking behavior and close social interactions with pair mates in Aotus may influence bacterial transmission and their glandular microbiomes. Chemical communication, including coordinated scent marking, may play a role in strengthening pair bonds, signaling pair status and/or in mate guarding in this socially monogamous primate.

Red foxes avoid apex predation without increasing fear

AbstractApex predators structure ecosystems by hunting mesopredators and herbivores. These trophic cascades are driven not only by the number of animals they kill, but also by how prey alter their behaviors to reduce risk. The different levels of risk navigated by prey has been likened to a “landscape of fear.” In Australia, dingoes are known to suppress red fox populations, driving a trophic cascade. However, most of what we know of this relationship comes from circumstances where predators are persecuted, which can affect their social and trophic interactions. Utilizing camera traps, we monitored fox behavior when accessing key resource points used by territorial dingoes, in a region where both predators are protected. We predicted that foxes would avoid and be more cautious in areas of high dingo activity. Indeed, foxes avoided directly encountering dingoes. However, contrary to our expectations, foxes were not more cautious or vigilant where dingo activity was high. In fact, fox activity and scent-marking rates increased where dingo scent-marking was concentrated. Further, foxes were increasingly confident with increasing levels of conspecific activity. Our results suggest that responses to the threat of predation are more complex than fear alone. In socially stable conditions, it is possible that prey may develop knowledge of their predators, facilitating avoidance, and reducing fear.

Does human infrastructure shape scent marking in a solitary felid?

Human development, such as construction of roads and trails, can affect several animal behaviour traits and is increasing worldwide. Despite the importance of scent marking for communication and social organization in many mammals, it is not clearly understood how scent marking is affected by human habitat modification. We used snow tracking data collected across six winters to study the effects of road infrastructure on the marking behaviour of Eurasian lynx (Lynx lynx) in a human-dominated landscape. We found that lynx marked at higher rates (11.2 urine sprays/km) on human routes (forest roads and logging trails) than when walking in natural habitat (5.8 sprays/km). This pattern was noted in both sexes, although males walked on human routes more frequently and scent marked more often than females. Compared to previous studies the mean marking rates we observed were the highest documented so far for wild felids. This may mainly be due to regular use of human routes (42% of all lynx movement) and the high scent marking rates found thereon. Human routes channelize lynx movement, and scent marking on these routes likely increases the probability of the chemical message being received by a conspecific; therefore human routes might enable more effective communication in territorial felids. Since most of the landscapes worldwide are covered by road networks and various human objects, the marking behaviour of wild mammals may be profoundly changed due to human-caused habitat modifications.

Is there a trade-off between scent marking and hunting behaviour in a stalking predator, the Eurasian lynx, Lynx lynx?

The costs of signalling are often expressed in terms of increased predation risk to the signaller; however, whether signalling predators also incur costs due to eavesdropping by prey and may attempt to reduce these costs is less well studied. In this study, we investigated whether there is a trade-off between intraspecific communication through scent marking and the risk of alerting prey in a wild population of a stalking predator, the Eurasian lynx. We followed lynx tracks in the snow and recorded scent marks and evidence of hunting behaviour along these tracks. Lynx preferred conspicuous objects for marking and increased scent marking rate when walking along linear structures, such as forest roads. This association was strongest when lynx were hunting, while there was only a weak correlation when no evidence of hunting could be detected. On tracks with evidence of hunting behaviour, lynx engaged less in scent marking. The relationship was most evident during the mating season, when lynx increased scent marking rates while they were not hunting. We further expected lynx to mark most in areas where they had recently hunted successfully, but time and distance to the last kill were not associated with scent marking rate. Our study supports the hypothesis that lynx face a trade-off between enhancing the detection probability of scent marks by conspecifics and avoiding eavesdropping by prey, and indicates that scent marking rate is influenced by several factors.

Scent marking in gerbils and its possible functions

Behaviors related to scent marking are compared and analyzed in males and females of four gerbil species (Meriones unguiculatus, M. tamariscinus, M. meridianus, and Psammomys obesus) observed in the wild and under semi-natural conditions. Scent-marking activity was found to vary in dependence on species, sex, age, reproductive conditions, social and territorial status of the individuals, and to show seasonal variation. The commonest patterns of scent marking are ventral rubbing and building up of ‘signal heaps’ with urine and feces. A close association between scent marking and social dominance was revealed in three species (M. unguiculatus, M. tamariscinus, M. meridianus). Spatial distribution of scent marks was found to be very uneven. Females marked most often the areas near burrow entrances, pathways and feeding sites. Males exhibited a higher rate of scent marking within home ranges of reproducing females. Inter-species differences in scent marking related to a species-specific space use systems and reproductive strategies were revealed. Results of the study partly support the scent-matching hypothesis and the status signaling hypothesis. However, these hypotheses do not predict spatial distribution of scent marks in male gerbils, and do not account for function of scent marking in young individuals as well as an increase of scent-marking activity induced by novelty factors. These findings are more consistent with the hypothesis of home range familiarization. Besides, scent marking in male gerbils could also function as a means of female monopolization.

Ontogenic patterns of scent marking in red foxes, Vulpes vulpes (Carnivora: Canidae)

Scent marking is widely recognised to have a crucial function in many species. Most research has focussed on adults and very little is known about scent marking patterns during juvenile development. Using video records of juvenile red fox Vulpes vulpes across 6 years, we tested whether scent marking rates varied with age or sex, or whether juveniles remaining on the natal territory (philopatry) marked more frequently than those that disappeared. Our data show that male juvenile red foxes scent marked more than females during early development, but rates rapidly declined as they aged. In contrast, females showed a significantly later and slower rate of decline. Within females, individuals that remained in the natal area had higher scent marking rates than those that disappeared, suggesting that scent marking has a role in social group affiliation within litters. These results demonstrate that scent marking plays an important role in juveniles, including their intra-litter social interactions.

Functional Context of Scent-Marking in Callithrix penicillata

Most mammals use olfactory signals for communicating identity, reproductive status and group-related dynamics. Callitrichids produce secretions as olfactory signals, the functional context of which is poorly known, especially in the wild. We analysed the functional significance of scent-marking in a free-ranging group of Callithrix penicillata in central Brazil. We recorded all occurrences of scent-marking according to context and conducted 10-min focal observations of adults and ‘scan sampling’ of the focal tree. We recorded the sex of scent-marking individuals and the location of occurrence within the home range, and found that there was no variation in rate of marking relative to either variable. However, females marked more along areas of contact with other groups in the home range and less in the core (non-contact) areas. Scent-marking in intergroup encounters, even when in agonistic contexts, was not associated with territorial disputes between groups. This suggests distinct functions for scent-marking for intra- versus intergroup communication. We also found a positive correlation between rate of scent-marking and height in the resource tree and between exudativory and air humidity.

The predation risks of interspecific eavesdropping: weasel-vole interactions

Competing species benefit from eavesdropping on each other's signals by learning about shared resources or predators. But conspicuous signals are also open to exploitation by eavesdropping predators and should also pose a threat to other sympatric prey species. In western Finland, sibling voles Microtus rossiameridionalis and field voles M. agrestis compete for food and space, and both species rely upon scent marks for intraspecific communication. Both vole species are prey to a range of terrestrial scent hunting predators such as least weasels, however, the competitively superior sibling voles are taken preferentially. We tested in large out-door enclosures whether field voles eavesdrop on the signals of its competitor, and whether they behave as though this eavesdropping carries a risk of predation. We presented field voles with scent marks from unknown conspecifics and sibling voles and measured their visitation, activity and scent marking behaviours at these scents under high (weasel present) and low (weasel absent) predation risk. Field voles readily visited both field and sibling vole scents under both high and low predation risk; however their activity at sibling vole scent marks declined significantly under increased predation risk. In contrast, predation risk did not affect field voles’ activity at conspecific scents. Thus, field voles were compelled to maintain eavesdropping on heterospecific scents under an increased risk of predation, however they compensated for this additional risk by reducing their activity at these risky scents. Scent marking rates declined significantly under high predation risk. Our results therefore reveal a hidden complexity in the use of social signals within multi-species assemblages that is clearly sensitive to the potential for increased predation risk. The predation risks of interspecific eavesdropping demonstrated here represents a significant generalisation of the concept of associational susceptibility.

Scent marking within and between groups of wild banded mongooses

Scent marking is commonly described as a territorial behaviour, and scent marks might deter potential intruders from entering occupied areas. Conspecific neighbours present both a reproductive and a territorial threat, thus, determining which, if any, of these threats shapes scent-marking behaviour is difficult. Banded mongooses Mungos mungo provide a rare clear separation between reproductive rivals (found within groups) and territorial rivals (neighbouring groups), because immigration into social groups is extremely rare, and mating occurs almost exclusively within groups. This situation offers an opportunity to assess the relative importance of territorial defence and intra-group competition for mates in shaping scent-marking behaviour. We combined detailed behavioural observations of scent marking, chemical analyses of scent composition and discrimination experiments in the field, and found little evidence for higher rates of scent marking in overlapping areas, a lack of group specificity of scents and a failure of individuals to discriminate between the scents of different groups. Although scent may fulfill some role in territorial demarcation and defence, these results suggest that scent marks and scent-marking patterns are also involved in communicating within social groups.

Interacting effects of predation risk and signal patchiness on activity and communication in house mice

1. Social signalling can be risky when signals are open to exploitation by eavesdropping predators. Unlike other signal modalities, olfactory signals cannot be 'switched off' in the presence of an eavesdropping predator, leaving receivers of scent signals at an increased risk of predation long after the signaller has moved on. Yet individuals of some olfactorily communicating species appear unwilling to forego the receiving of signals under an increased risk of predation. Foraging theory predicts that predation risk can operate at multiple spatial scales, however, such that prey behaviour should be sensitive to the broader olfactory environment beyond the risks of a single point source of odour. 2. Here, we use the house mouse Mus domesticus to test whether the spatial distribution and overall level of receiving activity varies with the spatial distribution of conspecific scent signals and the risks posed by an eavesdropping predator, the cat Felis catus. We assessed the mice's responses to these risks using overall visitation, activity and scent marking rates at conspecific scented locations (in clumped, random or regular distributions) and the surrounding matrix (non-scented) locations with and without a predator cue (cat urine). We then used univariate and bivariate spatial point pattern analyses to assess behavioural responses (activity) to both treatments across a range of spatial scales. 3. Visitation, activity and scent marking rates were not affected by the predator cue or the spatial distribution of scents. But these non-significant results masked a fine scale anti-predatory response. Mouse activity was significantly more clustered at small scales when in the presence of the predator cue; this response held across all spatial distribution treatments. Mice were also sensitive to the predation risks of clumped scents, and dispersed their activity at intermediate scales significantly more when exposed to the predator cue, than in the control scent treatment. 4. These results suggest that olfactorily communicating species use scale-sensitive anti-predatory behavioural changes to compensate for their increased risks of predation when receiving scent signals. We highlight the importance of examining a variety of scales when investigating predator-prey interactions, and discuss the implications of these findings for behaviourally responsive predators and prey.

Intergroup relations and home range use in Verreaux's sifaka (Propithecus verreauxi)

Relationships between neighboring groups feature prominently in socioecological theory, but few empirical studies have focused on the effects of neighbors on the behavior of primates. Interactions between neighboring groups are most conspicuous during intergroup encounters, but the likelihood of encounters with neighbors can also affect ranging and activity patterns indirectly, and, as a result, behavioral patterns in areas of exclusive use may differ from those in overlap areas of adjacent home ranges. The aim of this study was to examine intergroup relations and spatial variation in home range use in Verreaux's sifaka (Propithecus verreauxi) during the annual mating season. Based on 230 hr of focal animal data collected from ten members of five adjacent groups, we found that behavioral patterns and patterns of resource utilization were not markedly different between areas of exclusive use and overlap areas of adjacent home ranges. Group cohesion tended to be higher in overlap than in core areas, but the proportion of time spent resting and foraging did not differ between these two areas. However, dominant males exhibited a higher scent-marking rate in overlap areas. Observations during intergroup encounters revealed that chases between males occurred frequently, whereas fights involving physical contact were not observed. We also found that the intergroup dominance hierarchy depended on the relative group size or the number of males in each group, with groups of lower dominance rank exhibiting a tendency to sleep proportionally more often in their core areas. The results of this study suggest that in primate species with a moderate level of intergroup aggression, such as Verreaux's sifaka, the possibility of encountering neighbors in areas of home range overlap does not exert strong influence on their behavior and resource utilization.

Do cavity‐nesting primates reduce scent marking before retirement to avoid attracting predators to sleeping sites?

The largest population of endangered golden lion tamarins (Leontopithecus rosalia, GLTs) decreased from approximately 330 to 220 individuals between 1995 and 2000 due to a dramatic increase in predation at sleeping sites. We used behavioral data from eight social groups in this population to test two hypotheses: First, if GLTs attempt to mitigate the risk of predation at sleeping sites, they should reduce their rates of scent marking just prior to retirement. Second, if the benefits of scent marking prior to entering the sleeping site merit an increase in the rate of marking, then tamarins should increase their rate of pre-retirement scent marking during the breeding season, when such behavior would have its greatest impact on reproductive fitness. We used a generalized linear model (GLM) repeated-measures analysis to compare rates of daytime scent marking with rates of marking just prior to retirement for males and females. In addition, we compared scent marking prior to retiring in the nonbreeding season to marking rates before retirement in the breeding season for males and both sexes considered concurrently. Contrary to our expectations, GLTs significantly increased their rates of scent marking during the 30 min prior to entering their sleeping site-an observation driven by an increase in male (but not female) rates of marking. Rates of marking before entering the sleeping site were greater in the nonbreeding season compared to the breeding season, when both sexes were considered concomitantly and when males were evaluated alone. We conclude that GLTs do not attempt to minimize predation risk by decreasing scent marking in the period before they enter their sleeping site, and that tamarins do not scent mark at this time of day in order to transmit information about reproductive status or to control reproduction of subordinates. We speculate that scent marking in the 30 min prior to entering sleeping sites may serve to reduce predation risk by enabling tamarin groups to return quickly to favored sleeping sites in the evening when crepuscular predators are active.

RIVER OTTERS IN YELLOWSTONE LAKE DEPEND ON A DECLINING CUTTHROAT TROUT POPULATION

Introduced lake trout (Salvelinus namaycush) threaten the native cutthroat trout (Oncorhynchus clarki bouvieri) population in Yellowstone Lake, Yellowstone National Park, Wyoming. A reduction in availability of cutthroat trout may affect fish predators negatively, especially during spawning season. In this study we examined the importance of spawning cutthroat trout to a fish specialist—the North American river otter (Lontra canadensis). We measured scent-marking rates at 87 otter latrine sites on Yellowstone Lake and its tributary streams throughout the cutthroat trout spawning period and identified prey in scats collected at these sites. Based on fecal deposition rates, otters were more active on spawning streams and less active on Yellowstone Lake during the height of cutthroat trout spawning, with a return to elevated activity on the lake after spawning had ended. Cutthroat trout appeared to be the most common prey item in otter scats throughout the study, based on 515 samples identified to the family level and 110 samples analyzed to the species level. Overall, trout occurred in 72% and longnose suckers (Catostomus catostomus) in 43% of otter feces. Suckers were more prevalent than trout in otter scats only on tributary streams, toward the end of cutthroat trout spawning. Introduced lake trout, which inhabit deep water and are largely inaccessible to otters, occurred in less than 5% of otter scats. River otters in the Yellowstone Lake ecosystem appear to be relatively restricted in their diet and are heavily dependent on cutthroat trout. Our findings suggest that continued declines in the cutthroat trout population could negatively impact otters, potentially disrupting their role in linking aquatic and terrestrial ecosystems in and around Yellowstone Lake.

Sexual Dimorphism in Territorial Scent Marking by Adult Eurasian Beavers (Castor fiber)

Mammalian scent marking is often associated with territorial defense. However, males and females may demonstrate different activity patterns and play different roles. Female mammals nurture the young during lactation, while males purportedly perform other tasks more frequently, such as territorial maintenance and defense. This paper investigates the contribution made by mated pairs of adult males and females to territorial scent-marking in the obligate monogamous Eurasian beavers (Castor fiber). We hypothesized that both sexes should show territorial behavior, and predicted that they deposit a higher proportion of scent marks at borders. We also hypothesized that a sexual dimorphism exists due to reproductive constraints on the females, and predicted that males should invest significantly more in scent-marking behavior than females during summer. We obtained behavioral data by radio tracking six mated pairs of Eurasian beavers during spring and summer 2000-2001 on two rivers in southeastern Norway. Our results showed that both males and females clustered their scent marks near territorial borders, but males deposited a larger number of scent marks than females and spent more time at borders. Males were also found to have a higher scent marking rate and scent marks per night than females during summer, but not during spring. Overall, scent marks per night were higher in males than females. We conclude that both males and females Eurasian beavers carry out territorial behavior by scent marking, but males carry a larger part of the territorial defense during summer when females lactate. Our results are discussed in the light of the codefense hypothesis.

Scent marking strategies of new world primates

This paper reviews recent findings on scent-marking behavior in wild New World primates. These findings are used to examine three functional hypotheses: territoriality, regulation of social and reproductive dominance, and mating competition/mate attraction. Available data provide little evidence for a territorial function of scent-marking behavior in New World primates. Evidence for a function in the regulation of social and reproductive dominance is ambiguous. The patterns of scent marking found so far (i.e., rates of scent marking according to sex and social or reproductive status, and the spatial distribution of scent marks), and the sparse information on responses to scent marks are consistent with a hypothesized function in intrasexual competition and intersexual mate choice. Suggestions for future research are made.

Sex differences in scent marking in Propithecus edwardsi of Ranomafana National Park, Madagascar

In this study we compared the scent-marking rates of females with those of males. Specifically, we examined the ability of season, dominance status, and natal status to explain the frequency of scent marking in female sifakas living wild in Ranomafana National Park, Madagascar, and compared the results with those published for males [Pochron et al., American Journal of Primatology, in press]. We also sought to determine whether vulnerability to infanticide affects marking frequency in adults of either sex, and whether female reproductive status affects female marking behavior. We found that males marked at twice the rate of females, and like males, females in single-female groups marked at the highest rates. Dominant females and non-natal females marked at higher rates than did subordinate females and natal females, a pattern also seen in males. This suggests that scent marks may convey important information about status. Neither females nor males varied their marking frequency with the presence of vulnerable infants. Females did not alter marking frequency with reproductive state, and like males, they marked at higher rates in the period prior to the mating season than they did in the mating season itself. This implies that females may use scent marks more for intrasexual aggression or territoriality than for mate attraction.

Adult Male Scent-Marking in Lemur catta and Eulemur fulvus rufus

Scent-marking and olfactory communication are used extensively by prosimians and can provide spatial and temporal records of group movement and behavior. We compare rates of male scent-marking in relation to reproductive seasons, male dominance rank, and habitat use in two related prosimians: Lemur catta and Eulemur fulvus rufus. We collected scent-marking data on adult male Lemur catta at Beza-Mahafaly Reserve (dry forest), and on Eulemur fulvus rufus at Ranomafana National Park (rainforest), Madagascar. In Lemur, rates of overall scent-marking differed significantly by reproductive season, with higher rates occurring in mating and lactation/migration periods, whereas in Eulemur, reproductive season did not appear to affect scent-marking rates. Dominance rank of male Lemur catta did not affect rates of scent-marking. Among male Eulemur fulvus, dominance relations were not apparent; however, 2 of the 5 focal males scent-marked somewhat more frequently during the mating season and also experienced greater mating success. In Lemur catta, higher rates of scent-marking in the mating season may relate to indirect reproductive competition during a period of high aggression, while such mating competition was not as marked in Eulemur fulvus. Furthermore, higher rates of marking in resident male Lemur catta during male migration may correlate with vigilance toward immigrating males. Greater overall scent-marking rates in ring-tailed lemurs may relate to extensive intergroup home range overlap and no area of exclusive use, whereas the red-fronted lemur groups tended to forage in areas of their home range where little-to-no intergroup overlap occurred.

Artificially increasing scent mark rate increases urogenital gland size in mice Mus musculus

Male mice produce scent marks by depositing urine throughout their territory. The scent marks contain a number of pheromones and contain major urinary proteins (MUPs). Up to 1 g of these MUPs may be produced everyday. It is therefore likely that scent marking is costly and as such will impact on male fitness. In order to investigate the costs of scent marking, we conducted an experiment where scent marking rate was increased artificially, without changing the social environment (which may have an independent and different effect on both scent marking and physiology), or changing travelling time (such as would occur in a larger territory). Novel objects were introduced every other day (Replace); objects were introduced on day 1 and moved on alternate days (Move); or the mice were left undisturbed (Control). Introducing new objects daily caused a significant increase in scent marking rate, and an increase in the size of both the coagulating gland and the testes compared to the other two conditions. This is likely to be due to increased hormonal activity, which is known to affect these gland sizes. Interestingly, the preputial gland, which produces a number of pheromones, did not differ between the three conditions. There were no differences in male weight, growth rate or condition, indicating that the effect of an increase in scent marking of this magnitude does not have measurable fitness effects.