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.     

Variation in stimulus,seasonal context,and response to urine marks by captive Iberian wolves (<Emphasis Type="Italic">Canis lupus signatus</Emphasis>)

Urine marking behaviour was observed over 14 months in two captive pairs of Iberian wolves (Canis lupus signatus) by means of behaviour sampling. The study focused on the relative attractiveness of the different stimuli toward which urinations are directed, the seasonal context of urine marking, and the response by mates to sexually dimorphic action patterns. Kinds of marked substrates varied according to the posture used by wolves to urinate [raised-leg urination (RLU), flexed-leg urination (FLU), standing urination (STU), or squat urination (SQU)]. In RLUs and FLUs the wolves mainly used conspicuous substrates, whereas STUs and SQUs were carried out directly onto inconspicuous substrates. Wolves urinated on plants more than on other substrates, mostly selecting trees. A selection of trees according to their trunk diameter was also observed. The posts were chosen or avoided seemingly because of their specific characteristics such as size. In short, wolves marked with urine (RLUs and FLUs) those substrates that secured a greater effectiveness of marking. The mean duration of RLUs was lower than that of FLUs and SQUs. The rate of RLUs in males increased in winter (November–December) and in summer (July–August), while the rate of FLUs and SQUs in females and STUs in males did not increase during these periods. The wolves investigated RLUs more frequently than SQUs. The RLU display appears to be more ancestral than derived, similar in all species of canids and even in the Iberian wolf.     

Behavioral discrimination between circumgenital odor from peri-ovulatory dominant and anovulatory female common marmosets (Callithrix jacchus)

Social peer groups of callitrichid monkeys [marmosets and tamarins] exhibit intrasexual dominance hierarchies in captivity. This laboratory study employed two-choice behavioral discrimination bioassys to test the hypothesis that scent from female common marmosets contains chemical cues that permit discrimination between dominant females in the peri-ovulatory versus luteal phase of the ovarian cycle and females holding dominant versus subordinate status. When scent from only dominant females was presented, marmosets directed significantly greater amounts of investigatory behavior toward peri-ovulatory scent versus scent collected during the luteal phase of the ovarian cycle. Animals of both sexes demonstrated significant discriminatory behavior between scent deposited by dominant versus subordinate females, but only when the dominant female was in the peri-ovulatory phase of the ovarian cycle. Test animals directed equal amounts of investigative behavior toward scent from luteal-phase dominant females and subordinate females. Female test subjects deposited significantly more scent marks over presented scents than did male subjects, particularly when the scent had been donated by a peri-ovulatory female. Chemical odors specific to the peri-ovulatory and luteal phases of the ovarian cycle may play a role in mediating behavioral interactions among marmosets. Am. J. Primatol. 46:265–284, 1998. © 1998 Wiley-Liss, Inc.     

Scent‐Mark Identification and Scent‐Marking Behaviour in African Wild Dogs (Lycaon pictus)

Scent‐marking is common in mammals, but where signals are carried by urine and faeces, distinguishing between scent‐marking and mere elimination is problematic. To do so, we documented behaviours and context variables associated with urination and defecation in free‐ranging endangered African wild dogs (Lycaon pictus) and tested whether these were related to the responses of other dogs to deposits. We found that distinct postures were almost uniquely associated with deposits by dominant wild dogs, were more common during urination than defecation, and increased the likelihood that these deposits would be investigated by other wild dogs. The likelihood of investigation depended on the sex and dominance status of the depositor, the type of deposit and the substrate. Urine from dominant females was more likely to be investigated by other wild dogs than any other deposits, and deposits placed on vegetation were more likely to be investigated than those on bare ground. The likelihood that a deposit would be overmarked was affected by the deposit type and the sex and dominance status of the last depositor. Collectively, these results suggest that dominant wild dog urine is of greatest interest to other dogs. Our results show that some deposits by African wild dogs are not scent‐marks and that detailed observations of behaviours and context variables during elimination events can be used to distinguish deposits that are likely to be of communication value.     

Sociosexual development,pair bond formation,and mechanisms of fertility suppression in female cotton-top tamarins (Saguinus oedipus oedipus)

The effect of various social environments on sociosexual behavior was examined in six young female cotton-top tamarins (Saguinus oedipus oedipus) and in three established breeding females. Behavioral observations and hormonal samples were collected on young females while they were living with their families, when they were isolated from conspecifics, and after they were paired with an unrelated male. While living with the family, all females showed a suppression of fertility and low frequencies of sociosexual behavior. Following removal from the family, isolated females displayed an increase in rate of scent marking and an increase in hormonal levels. When young females were paired with males, they were exposed to scent secretions from their natal families, from an unfamilar family, and from a control for a total of 24 weeks. After pairing, hormonal levels increased dramatically, and ovarian cyclicity began. An increase in sociosexual behavior and elevated levels of scent marking accompanied this physiological change. Newly paired females had higher rates of affiliative behavior and scent marking than did established breeding females. However, both newly paired and established breeding males were more likely to initiate contact, grooming bouts, and social sniffing than were females. Time to first ovulation was later in females who were exposed to scent secretions from their natal families than it was in those females given a control for the first 8 weeks following pairing. No female conceived during exposure to scent secretions. However, once normal ovarian cycling had begun or a pregnancy was established, exposure to scent secretions had no effect. Thus, the social environment influences the fertility, sociosexual behavior, and pair bond formation of cotton-top tamarins. In addition, chemical stimuli found in the scent secretions produced by the natal family are most likely involved in reproductive suppression.     

Scent‐Marking Behaviour in a Pack of Free‐Ranging Domestic Dogs

Most mammals scent‐mark, and a variety of hypotheses have been put forward to explain this behaviour. Most of our knowledge about scent marking in domestic dogs comes from studies carried out on laboratory or companion dogs, while few studies have been carried out on free‐ranging dogs. Here, we explored the functional significance of different scent‐marking behavioural patterns in a pack of free‐ranging domestic dogs by testing two non‐exclusive hypotheses: the indirect territorial defence and the dominance/threat hypotheses. Through direct observation, we recorded the locations of dog scent marks (urination, defecation and ground scratching) and information regarding the identity and posture of the marking animal. We found evidence that markings are used by dogs to form a ‘property line’ and to threaten rivals during agonistic conflicts. Both males and females utilized scent marking to assert dominance and probably to relocate food or maintain possession over it. Raised‐leg urination and ground scratching probably play a role in olfactory and visual communication in both males and females. Urinations released by females, especially through flexed‐leg posture, may also convey information about their reproductive state. Finally, our observations suggest that defecation does not play an essential role in olfactory communication among free‐ranging dogs and that standing and squat postures are associated with normal excretion. Our results suggest that many of the proposed functions of marking behaviours are not mutually exclusive, and all should be explored through detailed field and laboratory studies.     

Scent marking in two western Amazonian populations of woolly monkeys (Lagothrix lagotricha)

We describe patterns of scent marking observed in two wild populations of lowland woolly monkeys that were the subjects of long-term studies in the westernmost portion of the Amazon basin. The woolly monkeys engaged primarily in two types of scent marking: chest rubbing and anogenital rubbing. In both study populations, males and females performed both types of scent marking, but males chest-rubbed more commonly than females, while females engaged in more anogenital rubbing. We evaluated two nonexclusive hypotheses for the function of scent marking by woolly monkeys: 1) that scent marking is used in sociosexual contexts, and 2) that scent marking is used to convey information about occupancy of or willingness to defend an area from conspecifics in other social groups. We found no association between the occurrence of scent-marking behavior and location within the home range, but did find that scent marking occurred more commonly than expected on days when copulations, mating solicitations, and intergroup encounters were observed. Additionally, mating activity and chest rubbing were highly correlated across the yearly cycle, even when the potentially confounding variable of ripe fruit availability was controlled for. In woolly monkeys, overt male-male competition is rare and female choice is an important part of the mating system. Our results are most consistent with the idea that scent marking plays a role in advertising male quality or competitive ability, and perhaps in coordinating mating activity.     

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.     

Quantitative data on training new world primates to urinate

This study assessed the effectiveness of operant conditioning in training three species of captive callitrichid primates (Leontopithecus rosalia, Callithrix geoffroyi, and Saguinus imperator) to urinate on demand. There were three goals to the study: 1) to develop a system for quantitatively assessing positive reinforcement training; 2) to ascertain whether or not positive reinforcement techniques can be used to train callitrichid monkeys to urinate on demand, and if so, how many training sessions are required; and 3) to determine the effect on urination behavior of the trainer entering the cage to collect a urine sample. Positive reinforcement with a continuous reinforcement schedule was used to capture a natural behavior: urination. Training sessions (30 min each) were conducted at dawn thrice weekly during five consecutive phases: habituation, control, training (animals were rewarded for urinating), maintenance (animals had reached a defined training criteria and continued to be rewarded for urinating), and collection (animals were rewarded for urinating, and the trainer entered the cage to collect the sample). The numbers of 30-min training sessions required to train the three monkey species (L. rosalia, C. geoffroyi, and S. imperator) were five, six, and eight, respectively. For the three species, the mean number of urinations per animal was significantly greater during the training, maintenance, and collection phases compared to the control phase. However, the three species differed significantly in the manner in which the rates of urination changed across the five phases. A higher proportion of subjects urinated during the training, maintenance, and collection phases compared to the control phase. Latency to first urination varied significantly across the five phases, with significantly reduced latencies to urinate during the training, maintenance, and collection phases compared to the control phase. The entry of the trainer into the cage to collect the urine sample did not appear to alter urination behavior. We demonstrate that operant conditioning techniques, which typically incur minimal cost, time investment, and disturbance, can be used to increase the quantity of urine samples collected for physiological analysis, the proportion of animals that urinate, and the speed of sample collection.     

The effects of gonadal hormones on scent marking and related behavior and morphology in female gray short-tailed opossums (Monodelphis domestica)

The hormonal control of scent marking and related behavior and morphology was examined in female gray short-tailed opossums. Females rarely scent marked when intact or following ovariectomy. Testosterone (T) but not estradiol (E) treatment stimulated chest marking while either hormone stimulated head marking in ovariectomized females tested alone. When the same females were tested with males, T-treated females showed little scent marking of any type; E-treated females showed hip marking in significantly more tests than females in the other treatment groups. Suprasternal scent glands (absent in intact females) and phalluses of females that received T were significantly larger than those of animals that received E or control animals. These findings are discussed with respect to similarities and differences between marsupial and eutherian females and between male and female gray opossums in the hormonal control of sexually dimorphic behavior and morphology.     

Female-Directed Aggression in Free-Ranging Ateles geoffroyi

In a study of the reproductive biology and behavior of black-handed spider monkeys (Ateles geoffroyi) on Barro Colorado Island, Panama, 107 instances of female directed male aggression (male attacks) occurred during >1,000 h of observation. Two hypotheses might explain this behavior. First, males may attack to induce defecation/urination by target females, from which they can gain reproductive information. Rates of defecation and urination did not increase subsequent to attacks; however, there was an association between attacks and investigative behaviors such as place sniffing. Alternatively, the attacks might be a form of sexual coercion. My results indicate that female reproductive state, as determined by fecal steroid metabolites (E1C and PdG), had an inconsistent effect on the frequency of attacks received. In addition, when females were in the peri-ovulatory period of the ovarian cycle they did not receive more attacks than at other times. Nor were attacks associated with observed copulations, suggesting that sexual coercion does not sufficiently explain the behavior. Some support is provided for the notion that the attacks allow males to dominate equally-sized females.     

Scent-marking behaviour by male prairie voles,Microtus ochrogaster,in response to the scent of opposite- and same-sex conspecifics

We conducted an experiment using the prairie vole (Microtus ochrogaster) to test predictions associated with the proposed functions of scent marking as a sexual attractant, in reproductive competition, and as a self-advertisement. We allowed an oestrous female, an anoestrous female, and an adult male to scent mark three portions of a clean substrate and then exposed a second male to this substrate for secondary marking. We did not support a sexual attraction hypothesis in that males did not place more scent marks in response to oestrous than anoestrous females. Similarly, we did not support a reproductive competition hypothesis in that males did not place more scent marks in response to marks of males than to those of females or bare substrate. Males did not overmark the scent of males or females and thus we did not support a scent-masking or scent-blending hypothesis. In that males deposited scent similarly in response to males, females, and on bare substrate, our results suggest that the frequency and placement of scent marks by males function primarily to advertise individual identity in an area.     

Do Male Veiled Chameleons,Chamaeleo calyptratus,Adjust their Courtship Displays in Response to Female Reproductive Status?

Variation in male courtship behavior may be due to inherent differences among males or may arise from males adjusting their courtship displays according to female responsiveness. Female veiled chameleons, Chamaeleo calyptratus , exhibit two distinctive suites of body coloration and behavior patterns that vary according to receptive and non-receptive stages of their reproductive cycle. We presented male chameleons with both receptive and non-receptive females, and recorded differences in their mating frequency, courtship intensity and courtship behavior patterns. As expected, males were more likely to court and attempt mating with receptive females. Although fewer males courted non-receptive females, their courtship displays were significantly longer than those directed towards receptive females. Males also adjusted the contents of their displays according to female reproductive condition. Certain behavior patterns were unique to courtship displays directed towards each class of females. Males exhibited the behavior pattern `head roll' only when paired with receptive females, and `chin rub' was displayed only during courtship of non-receptive females. We hypothesize that these differences in male courtship frequency, intensity and content reflect differences in female reproductive value. Although males may benefit from mating with both receptive and non-receptive females, the costs associated with courtship may depend on female responsiveness. Thus, males adjust their courtship tactics accordingly.     

Scent marking and mate choice in the prairie vole, Microtus ochrogaster

Scent marking is common among male and female rodents and might be used in male-male competition and as a mechanism for mate attraction. I tested the hypotheses that females would choose males based on their frequency and placement of scent marks, and that a female would advertise interest in a particular male by placing her scent marks on or near those of a preferred mating partner. In a series of experiments conducted with prairie voles, Microtus ochrogaster, females did not choose mates based on the frequency or placement of scent marks by males nor did they advertise their interest in a particular male through the frequency or placement of scent marks. The number of males chosen that scent-marked more than their opponents did not differ significantly between females exposed (11 of 15) and not exposed (10 of 15) to scents of males. Females exposed and not exposed to scents of males preferred seven of the same males that had scent-marked more than their opponents. When a third group of females was exposed to four times more scent of the less preferred than preferred males, they still chose the preferred males. Thus, the frequency and placement of scent marks by males were not used to assess males for mate choice nor did female prairie voles use scent to advertise their preference for a mating partner. In that scent marking is common in male and female mammals, scent quality might be more important than quantity in male-male competition and mate attraction.Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved .     

Fecundity and longevity of roaming dogs in Jaipur,India

Background

Estimates of demographic parameters, such as age-specific survival and fecundity, age at first pregnancy and litter size, are required for roaming dogs (i.e. dogs that are neither confined nor restricted) to assess the likely effect of proposed methods of population control. Data resulting from individual identification of dogs spayed as part of an Animal Birth Control (ABC) programme in Jaipur, India, are used to derive such parameters for the roaming dog population of that city.

Results

The percentage of females becoming pregnant in any given year was estimated by inspection of over 25,000 females caught for spaying from 1995 to 2006. The point estimate is 47.5% with a 95% confidence interval from 44% to 51%. Adult annual survival of spayed females was estimated by recapture of 62 spayed females from 2002 to 2006. The point estimate is 0.70 (95% confidence interval from 0.62 to 0.78), corresponding to an expected total lifespan of 3.8 years for a spayed female at one year old.

Conclusion

Recording the pregnancy status of dogs collected for spaying and individual marking of dogs released following spaying can provide estimates of some of the demographic parameters essential for predicting the future effectiveness of an ABC programme. Further, we suggest that recording the number and location of spayed and unspayed dogs encountered by the catching teams could be the most effective way to monitor the size and composition of the roaming dog population.

     

Effects of age, sex, and reproductive status on scent marking frequencies in the golden lion tamarin, Leontopithecus rosalia

This study reports on the longitudinal changes in scent marking frequencies of male and female lion tamarins in relation to age, reproductive status, and group composition. Juvenile males and females in family groups and subordinate males in trios (2 males, 1 female) scent marked infrequently. Juvenile females scent marked less than juvenile males and began scent marking later. Adult pair-bonded males and females scent marked with similar frequencies. Females showed scent marking increases during mid-pregnancy, but their scent marking decreased both before and again after parturition. Both males and females exhibited decreases in scent marking while transporting dependent infants.     

Meadow Voles (Microtus pennsylvanicus, Arvicolidae) Over-mark and Adjacent-mark the Scent Marks of Same-sex Conspecifics

Scent over-marking occurs when an animal deposits its scent mark on top of the scent mark of a conspecific; adjacent-marking occurs when an animal deposits its scent mark next to the scent mark of a conspecific. Given that male rodents usually scent mark more than females and that animals spend more time investigating the odor of the top-scent donor of an over-mark, I tested the following three hypotheses. First, male meadow voles deposit more scent marks than female meadow voles. Second, male meadow voles will deposit more over-marks and adjacent-marks in response to the scent marks of a same-sex conspecific than females would. Third, meadow voles spend more time investigating the odor of the second vole placed in the arena than that of the first vole placed in the arena. To test these hypotheses, two age-matched, like-sex conspecifics (first vole and second vole) were placed successively into an arena in which they were allowed to freely explore and scent mark for 15 min. The first hypothesis was not supported. The first and second vole, independently of sex, deposited a similar number of scent marks. The second hypothesis was also not supported by the data: more conspecific scent marks were over-marked by the second female than by the second male. The third hypothesis was supported by the data. After investigating a scented arena, males and females spent more time investigating the odor of the second vole than that of the first vole. Sex differences in scent-marking behaviors of meadow voles are unlike those reported for other species of rodents.     

Vestibular, vaginal and urethral relationships in spayed and intact normal dogs

The purpose of this study was to evaluate the urogenital anatomy in sound spayed and intact dogs. Fourteen intact and 12 spayed sound dogs had their lower urogenital tract evaluated with conventional vaginourethrography, computed tomography (CT) vaginourethrography and uroendoscopy under general anesthesia. Measurements for each of the three imaging modalities were made. Measurements included vaginal length, vaginal height, vaginal width, vestibule length, vestibule height, vestibule width, cingulum height, cingulum width, cingulum area, urethral length, urethral height, urethral width, angle of urethra to vestibule and angle of vagina to vestibule. Group comparisons were made between sound spayed and intact dogs. In general, most measurements were greater in intact dogs compared to spayed dogs regardless of imaging modality utilized. Group-weight interactions (P<0.01) were found on conventional vaginourethrograms (vaginal height and length and vestibule height, length and width) and CT vaginourethrograms (vaginal length, vestibule height, length and width, cingulum area, urethral width and angle of vagina to vestibule). A three-way interaction (P<0.01) for vaginal length, vaginal height and vestibule height was also identified. Vestibule length and height, cingulum height and urethral length were larger (P<0.01) with increasing dog weight and in intact dogs for both conventional and CT vaginourethrograms. No difference in vestibulovaginal ratio was observed between intact dogs and spayed dogs (P=0.0221). One of 14 intact dogs and 8 of 12 spayed dogs had vestibulovaginal ratios <0.33 on conventional vaginourethrograms. Anatomical differences in the lower urogenital tract between spayed and intact dogs emphasizes the need to establish normal reference ranges for spayed and intact dogs.     

Male aggression and the sexual behavior of Japanese monkeys

The factors that regulate male aggressive behavior towards females during the mating season and their relationship to the sexual behavior of Japanese monkeys were investigated. Observations were made on the Shiga A troop in Nagano Prefecture for 132 days during both the non-mating and mating seasons in three successive years. As a result, chasing towards females was observed in males older than 4 years. The frequencies of this type of behavior increased in the mating season. The chasing rank of the individual males did not correlate with the male dominance rank but with the tree-shaking rank. Chasing was directed not only at estrous females but also at non-estrous females. Pairs of both sexes in which chasing was observed, tended to have sexual interactions. Both hormonal and social factors should be considered in the regulation of chasing. The role of aggression in the formation of new male-female bonds in discussed.     

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.