Cheetah behaviour at scent-marking sites indicates differential use by sex and social rank

Scent-marking is an important form of communication for solitary species, as the information remains for the receiver after the sender has left the area. Individuals of different demographic or dominance classes (ranks) need to provide different information to conspecifics. Therefore, the use of scent-marking sites is of particular value for solitary species, but their functional role may vary between demographic classes. Using camera traps, we video-recorded the behaviour of cheetahs at scent-marking sites, assessing how individuals of different sex and social rank use the sites for different roles. We tested hypotheses that these sites are for territorial marking, reproductive signalling and information gathering. We found that females visit the sites infrequently, apparently to signal oestrous events, while males visit the sites frequently for territorial marking and information gathering. Among males, dominant individuals scent-marked the most and reacted to female visitation by vocalizing. In contrast, subordinate males, although also visiting the sites, were not recorded scent-marking nor vocalizing. These differences in the functional use of scent-marking sites among sex and social rank support the hypotheses that cheetah s use these scent-marking sites for both territorial and reproductive signalling, as well as to gather information.     

The role of scent marking in the social communication of wild golden lion tamarins, Leontopithecus rosalia

The role of scent marking in the social communication of mammals is widely variable. One reason for this variation is that the function of scent marking may vary with different ecological and social conditions. The purpose of this study was to test four nonexclusive hypotheses explaining the role of scent-marking frequency in different ecological and social contexts for wild golden lion tamarins. Relative to ecological contexts, we compared scent-marking frequency during seasons of abundant and scarce food resources. Relative to social contexts, we compared scent-marking frequency when groups were isolated and when groups were in the presence of neighbouring groups. We found that the tamarins used scent marking to mark the location of food resources. Additionally, males used scent marking to communicate intrasexual dominance within their groups, while females did not. Our results also indicate that alpha females increased their scent-marking frequency to communicate to members of other groups, while the presence of members of other groups did not elicit a similar response by alpha males. We did not find evidence for a territorial function of scent marking in golden lion tamarins. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour      

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.     

Variations in scent-marking behaviour of European badgers<Emphasis Type="Italic">Meles meles</Emphasis> in the vicinity of their setts

For European badgersMeles meles (Linnaeus, 1758), the importance of olfactory signals located at home-range borders in the context of territoriality has been widely accepted. Badgers, however, also scent mark far from their borders, often in the vicinity of their communal sett. Little is known about the significance of these marks in intraspecific communication. Here, we investigated the patterns of object-marking with subcaudal gland secretions close to the sett. Using remote-controlled, battery-powered infra-red video equipment, we recorded 442 incidences of object-marking between April 1996 and June 1997. The frequency of object marking varied significantly in relation to season, sex, age and reproductive status. In both sexes, relative object-marking rates were highest during the mating season, when individuals in reproductive condition marked significantly more often than non-reproductive animals. During the cubrearing season females marked at a significantly higher rate than males, and in both sexes adults scent-marked significantly more frequently than younger individuals. Approximately 30% of all scent-marks received an over-mark within 24 h of their deposit. In males over-marking behaviour was recorded only during the mating season, whereas females over-marked in all seasons at equal rates. Overall, our results suggest that in addition to their territorial functions, subcaudal scent-marks also serve as individual-specific advertisement signals directed at other group-members.     

Scent-marking in captive red pandas (Ailurus fulgens)

Three male and three female captive red pandas (Ailurus fulgens) were observed during and outside of the breeding season to examine changes in scent-marking behavior. Variation in scent-marking was analyzed with respect to site preferences, sex differences, onset of breeding, behavioral correlates, and environmental factors. Both males and females exhibited significant preferences for particular marking sites, those being prominent points in the enclosures. Males showed significantly higher scent-marking frequencies than females, except in one enclosure where increased marking behavior was associated with the presence of young. Sexual dimorphism in the pattern of marking was also observed, with males showing a bidirectional mark and females a unidirectional mark. Female marking was greater during the breeding season. Sniffing and number of waddles per mark were associated with scent-marking. Temperature change was not related to frequency of scent-marking. These results are compared with scent-marking behaviors in other species of carnivores and functional explanations are suggested.     

Age Affects Over-Marking of Opposite-Sex Scent Marks in Meadow Voles, Microtus pennsylvanicus

Models of age-related effects on behavior predict that among short-lived species younger adults are more attractive and attracted to opposite-sex conspecifics than are older adults, whereas the converse is predicted for long-lived species. Although most studies of age-related effects on behavior support these predictions, they are not supported by many studies of scent marking, a behavior used in mate attraction. Over-marking, a form of scent marking, is a tactic used by many terrestrial mammals to convey information about themselves to opposite-sex conspecifics. The present study tested the hypothesis that the age of meadow voles, Microtus pennsylvanicus ; a microtine rodent, affects their over- and scent-marking behaviors when they encounter the marks of opposite-sex conspecifics. Sex differences existed in the over-marking behavior of adult voles among the three different age groups that were tested. Male voles that were 5–7 and 10–12 mo olds over-marked a higher proportion of the marks of females than did 2–3 mo old male voles. Female voles that were 2–3, 5–7, and 10–12 mo old over-marked a similar number of marks deposited by male voles. Overall, the data were not consistent with models predicting the behavior of short-lived animals such as rodents when they encounter the opposite sex. The differences in over-marking displayed by older and younger adult male voles may be associated with life history tradeoffs, the likelihood that they will encounter sexually receptive females, and being selected as mates.     

Scent-marking behaviour of the honey badger, Mellivora capensis (Mustelidae), in the southern Kalahari

We investigated sexual and seasonal patterns in scent-marking behaviour of the honey badger, by direct observations of habituated individuals (five females, four adult males, two young males). Four categories of scent-marking behaviour were identified: (1) scent marking at latrines; (2) token urination in holes along the foraging path; (3) squat marking at single-use sites; and (4) functional excretion. Females and young males used all four types of scent marking, but adult males were not observed to use token urination. A strategy of hinterland scent marking was used, as was predicted from the large home ranges of both male and female honey badgers. There were significant sexual differences in marking rate: adult males primarily used latrines and adult females favoured token urination. Latrine scent marking in adult male honey badgers provides support for the ‘scent-matching’ hypothesis. Females visited latrines when they were in oestrus. However, the low level of marking activity during a visit and the intensive smelling suggested a scent-matching function rather than reproductive advertisement. Token urination appeared to be related to the maintenance of spatiotemporal separation in females, although we also observed token urination in young males. While the placement of urine in foraging holes and its relation with successful digging attempts offer some support for the foraging efficiency hypothesis, we consider this unlikely, because we did not observe it in adult males and there was no seasonal pattern. Squat marking occurred under a wide range of conditions in both males and females and may be related to marking valuable resources. It is likely that scent marking in honey badgers has many functions.     

The effect of pair bonding in Cabrera vole’s scent marking

The Cabrera vole (Microtus cabrerae) is a rare rodent living in patchy grassy areas of the Iberian Peninsula where unpaired individuals of both sexes use scent marking primarily to increase their mate-finding likelihood. Cabrera voles establish long-term pair bonds with opposite-sex conspecifics constituting a breeding pair, which is expected to reduce the efforts in searching for a new mate. Under such circumstances, scent marking as a strategy to increase mate-finding likelihood became useless. Accordingly, we hypothesise that pair bonded Cabrera voles suppress mate-finding scent marking to reduce energetic costs and predation risk. To test this hypothesis, we compared scent-marking behaviour towards a clean substrate, in both paired and non-paired voles. No differences were found in the scent marks’ type and the amount of marks placed by voles in both conditions. We also analysed the scent-marking behaviour of both sex pair bonded voles when exposed simultaneously to a clean substrate, a substrate pre-marked by males and a substrate pre-marked by females. We found no significant differences in scent-marks (urine-marked area and number of faecal boli) across the three types of substrate types. In accordance with our prediction, these results suggest that pair bonded Cabrera voles did not use scent marking for mate finding, thus providing further support to the existence of a monogamous mating strategy. Furthermore, our results fail to support the use of scent marking for territorial defence purposes.     

Marking behaviour in two lemur species (L. fulvus and L. macaco): relation to social status, reproduction, aggression and environmental change.

The influence of social status, reproductive and agonistic contexts and environmental change on scent-marking and allomarking rates were studied in captive groups of Lemur fulvus and Lemur macaco. No evidence of female social dominance over males appeared in either species. In L. fulvus, intrasexual relationships were characterized by the dominance of one adult male over another and by the existence of female affiliative association ('central' vs. 'peripheral' females). In L. macaco, no intrasexual dominance relationships were apparent except for a brief ostracism of one female. In both species, (1) intrasexual differences in social status were related to differences in marking rates, (2) no direct relation appeared between marking rates and aggression or reproduction, (3) allomarking was not differentially directed towards specific individuals and (4) physical environmental factors clearly influenced scent-marking and allomarking rates. These results are discussed in relation to the possible functions of marking behaviour in intragroup relations.     

Scent-marking in the tamarin, Saguinus oedipus: Sex differences and ontogeny

Many mammalian species which exhibit scent-marking behaviour show a pronounced sexual dimorphism in marking behaviour and scent gland morphology. However, several species of marmosets and tamarins do not show dimorphism in these traits. We examined sex differences in scent-marking structures and behaviour in cotton-top tamarins (Saguinus o. oedipus; Primates: Callitrichidae). While body size and weight were virtually identical in adult males and females, there were pronounced sex differences in scent gland size and in rates of marking behaviour. Females possessed larger anogenital and suprapubic glands and showed 10-fold higher rates of anogenital marking and slightly higher rates of suprapubic marking than males. Observations on the development of anogenital marking revealed a lack of dimorphism during the first 2 years. Adult rates of anogenital marking in females were only observed in females housed separately from their natal family group. The onset of adult levels of marking behaviour corresponds with the adoption of the role of a breeding adult female, suggesting that anogenital marking plays a role in sexual communication.     

Scent-marking in free-ranging coyotes, Canis latrans

From November 1977 to March 1978 we recorded the scent-marking behaviour of free-ranging coyotes in southern Québec. Fourteen animals were marked by foot-pad ablation, which made it possible to recognize individuals' tracks in the snow and therefore to relate scent-marking with social organization. We observed 949 scent-marks. The identity of the animals was known for 50% of the tracking distances. Animals living in territorial groups (pairs and pairs with juveniles) marked more than twice as much as solitary non-territorial individuals. In addition the former used stronger forms of marking significantly more often than the latter.     

Scent-marking and intrasexual competition in a cooperative carnivore with low reproductive skew

Most mammals scent-mark and a variety of hypotheses have been put forward to explain this behaviour. Differences in the main function of scent-marking between species are likely to be related to differences in social systems. Here, we investigate the functions of scent-marking in a cooperatively breeding carnivore. In the banded mongoose ( Mungos mungo ), individuals of both sexes commonly breed in their natal group and reproductive skew within groups is low. Using experimental scent-mark presentations, we tested predictions of the intrasexual competition, self-advertisement to potential mates and dominance assertion hypotheses. Both males and females responded more intensely to scent marks of same-sexed than of opposite-sexed individuals. Dominant individuals counter-marked more than subordinate ones and males showed higher counter-marking rates than females, but only marginally so. During oestrus, responses to scent marks were increased by both sexes. Our findings strongly indicate that scent-marking in the banded mongoose primarily serves a purpose in intrasexual competition both between and within groups. Unlike in other social herpestids and some solitary rodents, we found little evidence for self-advertisement. We suggest that the peculiar social system of the banded mongoose results in self-advertisement losing importance in this species, shifting the main function of scent-marking to intrasexual competition.     

Seasonal and sex differences in urine marking rates of wild red foxes Vulpes vulpes

Understanding the role of urine marking in the territorial systems of wild mammals can be difficult, especially for nocturnal cryptic species. Even for common species, such as the red fox Vulpes vulpes, a comprehensive analysis of seasonal and sex differences has not been carried out. Using 6 years of infra-red video monitoring, we compared marking rates between months and between sexes. Urine marking was significantly lower during summer (June–August). Males urine marked significantly more frequently than females during late summer and autumn, but not winter. Males marked more frequently than females also during March. There was no increase during the breeding season for either sex. Our results correlate with previous partial data but demonstrate how urine marking rates vary across the year. They also further support the greater role of males in fox territorial maintenance. Urine marking is lowest during summer when territorial intrusions are least, whilst the higher male urine marking rate in March reflects the period when females are denning. Overall, our results provide the first comprehensive analysis of red fox urine marking rates, contributing to a greater understanding of territoriality and olfactory communication.     

Scent-marking behaviour in response to conspecific odours by the rat, Rattus norvegicus

The scent-marking behaviour of male and female rats in response to conspecific odours was investigated in two experiments. The results of the first experiment indicated that exposure to conspecific odours generally led to an increase in marking rates. However, no sex differences were found, and large differences in marking responses according to the sex of the odour donor were only found for dioestrous females. These results do not suggest that the primary function of scent-marking in this species is sexual. In the second experiment, odour sources differed not only in the sex of the donor, but also in their relative familiarity to the animal being tested. This experiment showed that males marked more in response to odours from familiar (non-related) females. Females did not differ significantly on measures of marking but did display greater interest in odours from unfamiliar males. The results are discussed in relation to possible functions of marking in this species.     

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.     

An observational study of scent-marking in coyotes, Canis latrans

Urination and defaecation patterns of free-ranging coyotes (Canis latrans) were studied in the Grand Teton National Park, Jackson, Wyoming, for two years. The vast majority of urinations by adult males and females were involved in ‘marking’, and differentiating between ‘marking’ and ‘elimination’ may not be necessary. Our results may be summarized as follows: (1) raised-leg urinations (RLU) performed by males were most frequently used in marking. (2) Females marked throughout the year using the squat (SQU) posture. (3) Snow tracking and reading snow sign resulted in a gross underestimate of the relative frequency of SQU's and a large overestimate in the relative frequency of defaecations (DEF) when compared to results obtained by direct observation. (4) There was sexual dimorphism for the contexts in which marking occurred. Overall, marking by males was associated with courtship and mating, with travelling, and with aggression. Marking by females was associated with the acquisition and possession of food and with the denning season. (5) Marking rates per coyote increased in groups larger than two animals. (6) RLU marking rates were greatest in areas of high intrusion when compared to denning areas and areas in which non-group members infrequently tresscent odours are important in orienting individuals in space but do not represent in and of themselves barriers to movement.     

Scent marking in a territorial African antelope: I. The maintenance of borders between male oribi

Scent marking is ubiquitous among the dwarf antelope and gazelles of Africa, but its function has been the subject of debate. This study examined preorbital gland scent marking in the oribi, Ourebia ourebi, a territorial African antelope. Several hypotheses for the function of scent marking by territorial antelope were tested with observational data. Of these, the hypotheses that scent marking is driven by intrasexual competition between neighbouring males, and that marks serve as an honest advertisement of a male's ability to defend his territory from rivals, were supported best. Thirty-three territorial male oribi on 23 territories marked most at borders shared with other territorial males, and territorial males marked more often at borders shared with multimale groups than at borders shared with a single male. This suggests that males perceived neighbouring male groups as a greater threat to territory ownership than neighbouring males that defended their territories without the aid of adult subordinates. Marking rate was unrelated to territory size or the number of females on adjacent territories, but males with many male neighbours marked at higher rates than those with fewer male neighbours. These results suggest that the presence of male neighbours has a greater effect on the scent marking behaviour of territorial antelope than has been considered previously. Copyright 1999 The Association for the Study of Animal Behaviour.     

Sex difference in the communicatory significance of localized defecation sites in Arabian gazelles (Gazella arabica)

Mammalian scent marking in localized defecation sites (latrines) has often been interpreted in the context of (male) territory defense. However, latrines could have different functions in males and females, especially where territorial males monopolize groups of females with stable social alliances and pronounced home range overlap. We investigated the communicatory significance of latrines in wild Arabian gazelles (Gazella arabica) and assessed the spatial distribution of latrines within home ranges. Latrine density and utilization was highest in the center of female group home ranges, and less frequent in peripheral home range sections, pointing towards communication within groups rather than towards territoriality. When considering male home ranges, latrine densities and utilization were higher in non-overlap zones, contradicting a territorial function. This pattern appears to be caused by more females than territorial males per given area establishing latrines. A subsequent survey of latrine utilization, based on camera trapping, suggests that males use latrines for territory defense: males visited latrines in overlap zones disproportionally more often than females, and successions of two males prevailed. Our study thus highlights that male territorial marking can be masked when males and females use the same marking system for different purposes.     

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.     

Scent marking in sifaka: no one function explains it all

Various hypotheses to explain possible functions of scent-marking have been put forth and basically fall into five categories: territorial demarcation, ownership of resources, mate attraction, noncombative fighting, and self-advertisement. Verreaux's sifaka (Propithecus verreauxi verreauxi), like many prosimians, use scent-marking as one form of communication. The goal of this study was to determine the function of scent-marking in sifaka. All-occurrence of scent-marks, scent-mark context, and scent-mark style were collected on 23 sifaka in the Kirindy Forest of western Madagascar for 7 months (September 2001-March 2002). Scent-mark rates were collected using continuous focal animal sampling from November 2000-March 2002. Home range data were collected using monthly censuses and instantaneous focal sampling throughout the 17 months. Scent-marking behavior was exhibited almost exclusively by adults. Scent-mark rates were not related to the number of resident adult males, number of resident adult females, rank, or group size. The majority of scent-marks occurred in the perimeter of the home range; however, less than a quarter of the home range was used by only a single group. Sifaka did not preferentially mark food trees, mark during the mating season, or mark during intergroup encounters. The results from this study indicate that none of the five hypothesized functions best explains all scent-marking in Verreaux's sifaka. Rather, scent-marking may serve different functions for different individuals.