Social behaviour of captive bush dogs (Speothos venaticus)

The bush dog, Speothos venaticus , is a rare forest-dwelling South American canid. Social behaviour of a captive pack of three adult males and three adult females was observed over four years in a large enclosure. The dogs appeared to be compulsively sociable, sleeping in close physical contact, travelling together in single file, and feeding communally with minimal aggression. An alpha pair was dominant, but there was no clear hierarchy within the pack as a whole. There were, however, separate male and female hierarchies. The pack engaged in various communal behaviours, including greeting ceremonies in which individuals mutually submitted to one another. All dogs urine marked frequently, both sexes adopting postures to direct urine upwards and to soak their fur in odour. Only the alpha female bred successfully, and all other group members carried and guarded the young. These observations are discussed in the contexts of canid social behaviour and the breeding in captivity of this endangered species.     

Female preference for males that have exclusively marked or invaded territories depends on male presence and its identity in the subterranean rodent Ctenomys talarum

Territorial scent‐marking provides chemical records of male competitive interactions that are available to females, who gain valuable information to assess and identify best quality partners. In this context, the solitary subterranean rodent tuco‐tuco (Ctenomys talarum) offers excellent possibilities to evaluate the effects of male exclusive scent‐marking of territories on female assessment. For evaluation, we used wild caught individuals of C. talarum, manipulated their scent marks within the territories in captive conditions and staged preference tests where females were able to choose between exclusive and invaded territories. The evaluation was performed in two scenarios considering the identity of the intruder scent mark: territories invaded by a strange male and territories invaded by a neighbour male. Females investigated the chemical cues deposited on the substrate of the exclusively marked territory more frequently. Next, females displayed equal interest to scent samples of both males presented in a Y‐maze. Finally, when females could gain access to both individually isolated males and their scent‐marked territories, they spent more time within invaded territories despite they visited them with the same frequency. Moreover, females tried to get in contact by scratching the mesh of the owner of the invaded territory more frequently. We found that females of C. talarum evaluate the homogeneity (exclusiveness) of scent marks within a male territory and then show preferences in relation to the identity of the intruder's scent –whether strange or neighbour.     

Scent-Marking of Giant Otter in the Southern Pantanal, Brazil

Giant otters live in social groups, consisting of a mating pair and one or two litters. Groups are territorial and mark their territories often with scent-marks. Our objectives were to evaluate the frequencies of marking and over-marking according to the social status of the individuals and to define the different postures used during the marking. We observed four groups, totaling 25 individuals (five alpha males, four alpha females, seven adult females, one adult male and eight juveniles) with group size ranging between four and 13 individuals. The study was conducted between July 2006 and July 2007 in the Vermelho River and in a stretch of the Miranda River, in the Southern Pantanal. We observed the groups for a total of 2006 min and recorded 95 events of marking totaling 84.9 min. Time spent marking varied between groups and ranged from 4.3 to 44.7 min. The alpha males marked more frequently (62% of marking events, 55 min) than the alpha females (17% of marking events, 13.6 min). Of the 59 events of scent-marking by the alpha males, 32 over-marked the marks of other individuals from the group. Of the 16 events of scent-marking of the alpha females, five over-marked that of other females from the same group. When scent-marking, alpha males used the 'stepping' posture most frequently (63%), then 'fore-paw rubbing' (24%), 'latrine use' (7%), and 'body rubbing' (6%). Alpha females used the 'stepping' posture most frequently (65%), then 'latrine use' (19%) and 'fore-paw rubbing' (12%), with only one event of 'body rubbing' observed during marking. Subordinate females used the 'stepping' posture (76%) and 'latrine use' (24%) during marking. Scent-marking can play many roles in mammals and for giant otters, and the main roles appear to be communication of social and sexual status and territorial defense.     

A preliminary study of olfactory behavior of captiveLemur coronatus during the breeding season

The olfactory behavior of a captive population of crowned lemurs (Lemur coronatus)was studied during their annual breeding season in order to investigate its possible role in the fine-tuning of mating synchrony after photoperiodic initiation of reproductive activity. The frequencies of five stereotyped olfactory behavior patterns were recorded in four male-female groups during 315 1-hr observation sessions between October and March. The mean total scent-marking frequency of all males was positively correlated with their mean testicular size. Male anogenital-, head-, and allomarking frequencies decreased during the breeding season, whereas hand-marking frequencies remained constant. Male head-marking increased significantly in the 5 days preceding female vaginal estrus. Furthermore, males responded to many female scents by olfactory investigation and/or overmarking, whereas females never showed an observable behavioral response to male scent marks. The marking frequencies of two females did not change significantly during the breeding season, while those of two others decreased significantly. These results suggest qualitative changes of female scents during estrus and a less important role of male scents in intersexual communication in the context of reproduction than previously thought.     

Scent marking as resource defense by female Lemur catta

Because ring-tailed lemurs (Lemur catta) are a female-dominant, female-philopatric species in which the females provide the majority of parental care and troop defense, resource defense is a possible function of female lemur scent marking. To test this hypothesis, I conducted three studies. First, I presented captive, individually housed females with a series of samples of female scent, each from a different female, to determine whether they would respond to those samples and discriminate between them. Second, I reanalyzed data from a focal animal study of four females in two adjacent troops in Berenty Reserve, Madagascar, to determine female marking rates before, during, and after the mating season, and to clarify the relationship among positions of feeding, intertroop defense, and scent marking. The third study was based on ad libitum observations of the sniffing and marking behavior of a troop in Berenty Reserve during a year when they traveled far out of their home range. The females in study 1 investigated female scent samples but provided no evidence that they discriminated between them. In study 2 the wild females marked throughout the study and did not limit their marking to the mating season. They deposited significantly more of their marks in a zone of confrontation with adjacent troops, where they also did the majority of their feeding, and they increased their rate of marking during agonistic intertroop confrontations. The females determined the positions of their scent marks and deposited the first mark in the majority of countermarking sequences. When the females traveled out of their defended range in study 3, they significantly decreased their rate of marking and increased their rate of sniffing spots but not marking them. All evidence gathered so far supports the hypothesis that one function of female ring-tailed lemur scent marking is to provide intergroup information that is then used to reinforce the border of the defended resource.     

Urinary behavior of female domestic dogs (Canis familiaris): influence of reproductive status,location, and age

The urinary behavior of adult domestic dogs (Canis familiaris) is sexually dimorphic with respect to the posture (males lift a leg and females squat), frequency of urination (males urinate more frequently than females), and tendency to direct urine at specific objects in the environment (males are more likely than females to direct their urine). Such behavioral differences have led to the belief that urination functions largely, or exclusively, in elimination in female dogs, while having the additional function of scent marking in male dogs. In this study, we observed urinary behavior of six spayed and six non-estrous intact female Jack Russell Terriers during walks on and off their home area. The females ranged in age from 0.4 to 11.2 years. Frequency of urination was positively correlated with age, and females four or more years old directed the majority of their urinations at objects in the environment. Overall, females urinated more frequently and directed more of their urinations when walked off their home area than when walked within their home area. Spayed females were more likely than non-estrous intact females to ground-scratch following defecation; we detected a similar trend for ground-scratching after urination. There was, however, considerable variation among spayed females in the tendency to display ground-scratching behavior. Overall, the most common posture displayed by females while urinating was the squat-raise. Other postures, in order of their frequency of occurrence included squat, arch-raise, combination, and handstand. Females used the squat-raise and arch-raise postures more when off their home area than when on their home area. Overall, there was substantial individual variation among females in the postures used while urinating. Our data indicate that female urinary behavior varies with location and reproductive status, and that substantial individual differences exist among females for some patterns of behavior. Additionally, the large percentages of directed urinations by spayed (60.8%) and non-estrous intact females (56.7%) in our study suggest that urination in female dogs does not function solely in elimination, but that it also has a significant role in scent marking, even when females are not in estrus.     

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 .     

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.     

Olfactory communication and counter-marking in brown brocket deer <Emphasis Type="Italic">Mazama gouazoubira</Emphasis>

Scent marking by defecation and urination in numerous small latrines may be related to resource defence in brown brocket deer (Mazama gouazoubira). Both males and females seem to be territorial, and both contribute to latrines where their ranges overlap. Latrines could thus potentially function as centres of information exchange and intrasexual competition. Counter-marking occurs when animals respond to invaders' marks with a greater number of marks. The objectives of this experimental study were to determine whether brown brocket deer distinguish dung of presumed invaders from their own and whether they counter-mark such faecal deposits. Two samples of dung (from unknown males or females and the experimental animal) were introduced near the latrines of 21 captive deer (13 males and eight females), and we observed their responses, including investigative (sniffing) and marking (urination and defecation) behaviours. Males investigated the introduced dung and their own latrine significantly more when the dung was from an unknown male than when it was their own. Females investigated unknown female dung significantly more than their own. Males counter-marked the introduced dung and their latrine significantly more when the dung was from an unknown male than when it was their own. Males marked with a shorter latency and at a greater frequency than females. Our data indicated that males counter-mark most intensively dung from male ‘intruders’ which may be related to intrasexual competition and resource defence. Females showed a non-significant tendency to counter-mark same sex intruders.     

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.     

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.     

Determinants of Paternity Success in a Group of Captive Vervet Monkeys (<Emphasis Type="BoldItalic">Chlorocebus aethiops sabaeus</Emphasis>)

Paternity success of high-ranking primate males is affected by the number of males and the number of fertile females and their cycle synchrony. Female vervets in the wild show strong reproductive seasonality and do not advertise the ovulatory period with conspicuous signals or behavior. Because this makes it difficult for males to monopolize fertile females, it can be expected that male reproductive skew in this species is lower than in other cercopithecines living in multimale groups that advertise the ovulatory period. We assessed male reproductive success in a captive vervet group, initially consisting of 4 males and 12 unrelated females. Besides a general low reproductive skew, we predicted paternity success of the alpha males to be dependent on the overlap of synchronously fertile females, month into alpha male tenure, and housing conditions (the subjects were kept in a large park but had to be locked in a small indoor compartment during the winter months). Further, because the number of males reaching their prime increased over time, we predicted a decrease in reproductive success of the alpha male with increasing tenure length of the alpha male. To assess this we collected genetic samples from 27 of 32 offspring born into the group during 3 yr, using 7 polymorphic human microsatellite markers. Contrary to our predictions, reproductive success of the males occupying the alpha position during the study was among the highest found in cercopithecines, with the alpha males siring 78% of all offspring. The degree of overlapping fertile periods did not affect paternity success, nor did paternity success of the alpha male differ between the 2 housing conditions. Only the alpha male’s tenure length had an effect. Alpha male reproductive success decreased over time, which we attribute to either a shift in female mate choice toward younger males or to increased competition among the males in the later stage of the alpha male’s tenure. This study demonstrates the importance of assessing reproductive success of males throughout >1 season and should spawn more research into the role of female choice and postcopulatory mechanisms in this species.     

Correction to: The environment matters: season and female contact affect the behavior of captive Addax nasomaculatus male antelope

The addax antelope (Addax nasomaculatus) is a species under serious threat of extinction, as it is more abundant in captivity than in the wild. However, little is known about its basic biology. The aims of this study were to determine how locomotor, feeding, aggressive, marking, and sexual behavior of male addax allocated in all-male groups vary with season and with female contact (i.e., biostimulation). The study was conducted in captive conditions, in two groups of adult males: one with no-physical contact with females, aside from visual and olfactory interactions (CF group, n = 4), and another group completely isolated from females (IF group, n = 4). The frequency of behaviors was recorded during the daytime, 4 days per season (total time of observation = 256 h). Lying, standing, walking, aggressive, marking, grazing, and ruminating behaviors as well as water and supplement consumptions varied with season (all p < 0.05). The lying, walking, marking, grazing, and ruminating behaviors were more frequently observed for CF than IF males (all p < 0.05). Also, all behaviors, except for marking, varied with the interaction between the group and seasons (all p < 0.05). Sexual behavior was extremely scarce, so it was not possible to analyze how it varied with seasons and the group. The present study suggests that management program and housing conditions, especially in ex situ breeding plans, should consider the influence of the season and the sociosexual context on the behavior of addax males.

     

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

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

Female sexual behavior in the absence of male-male competition in captive Japanese macaques (Macaca fuscata)

A captive group of Japanese macaques (Macaca fuscata). consisting of one adult male and three adult females, was observed for 88 hr during the 1988 mating season. We focused on the group's sexual activities because understanding how social dynamics affect reproductive behavior may enhance our ability to establish self-sustaining captive populations. The adult male exhibited distinct preferences in copulation partners, although all females were receptive and cycled during the study period. The dominant female participated in the most copulations and successfully harassed and disrupted copulations between the male and the other females. The alpha female, therefore, actively constrained the formation of mating pairs. Further, the male did not consort with his year-round female grooming partner. This female, the least dominant member of the group, engaged in the most autosexual and homosexual behavior. Birth season data show that only the dominant female bore offspring. This study emphasizes the influence of social dynamics on a group's reproductive potential and suggests an alternative means by which females can influence consort formation.     

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.     

Behavior of giant pandas (Ailuropoda melanoleuca) in captive conditions: Gender differences and enclosure effects

The behavior of giant pandas (Ailuropoda melanoleuca) was studied under captive conditions. Both male and female pandas spent similar amounts of time engaged in eating and locomotion. Males performed anogenital‐marking more but rested less than females, which suggests a sexually dimorphic pattern of behavior. Furthermore, females housed in the seminatural environment spent significantly less time engaged in stereotyped behavior than did females housed in the traditional enclosure, indicating that an enclosure environment affects the behavior of giant pandas. These data illustrate the importance of careful management and facility design for captive giant pandas. Zoo Biol 22:77–82, 2003. © 2003 Wiley‐Liss, Inc.     

Discrimination between males and females by scent in Lemur fulvus.

Two captive male Lemur fulvus were presented with scents of conspecific males and females, outside the mating season. Both subjects sniffed male scents significantly more than female scents. Male and female scents did not elicit significantly different amounts of scent marking. None of the responses of either subject suggested any discrimination of sub-species by scent.     

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

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

Mating relationships and breeding suppression in the dwarf mongoose

Dwarf mongooses live in packs containing a dominant breeding pair. The alpha female produces litters at regular intervals, usually three times per year. Other sexually mature females come into oestrus in synchrony with the alpha female and occasionally become pregnant but may not raise their offspring. Some females which had not been visibly pregnant nursed the young of other females. Early in the alpha female's oestrous cycle the alpha male maintains proximity and copulates with her exclusively, attacking any other adult males which approach. Later he also copulates with other adult females and the beta male mates with the alpha female. The alpha pair are likely to be the parents of the great majority of young born in the pack.