Article

A Reassessment of the Function of Scent Marking in Territories

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Abstract

The energetic costs and the risk of injury in agonistic encounters can be reduced by prior assessment of opponents: it will generally pay low quality animals to avoid combat with one of high quality. Following this principle it is suggested that territory owners scent mark their territories to provide intruders with a means of assessment. When the odour of a competitor, or of a mark it is seen to have made, matches that of scent marks encountered in the vicinity, then the competitor is probably the territory owner. Since owners are generally high quality animals, and assuming they have more to gain by retaining a territory than an intruder has in taking it over, it will pay the owner to escalate and the intruder to give up early. The advantage to owners in marking may thus be that by allowing themselves to be identified they reduce the costs of territory defence. Published information on the behaviour of territory owners and intruders is consistent with predictions from this hypothesis. The hypothesis offers an explanation for a number of poorly understood behaviours including ‘self-anointing’ and scent marking during agonistic encounters.

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... Such behavior is referred to as scent marking or territory marking. A variety of functional explanations has been proposed for scent marking in mammals including rodents (reviewed by Gosling, 1982 and Roberts, 2007 ). According to the scentfence hypothesis, scent marks are like a fence around the periphery of the territory to keep intruders out (Hediger, 1949; Müller-Schwarze & Heckman, 1980). ...
... There is also a generally accepted idea that scent marking is a means of olfactory communication. In addition, scent marking is proposed to be a means by which individuals assess the competitive ability of opponents, and resource holders appear to mark to help establish and maintain their status (Gosling, 1982Gosling, , 1990 Gosling & McKay, 1990; Gosling & Roberts, 2001a , b). Roberts (2007) asserted that scent marking is a fundamental component of territorial behavior and of advertising dominance status within social hierarchies. ...
... The main goal of the study is an interspecific comparative analysis of different patterns of scent marking, its seasonal variability and functions in relation to species-specific space use systems. It was not my intention to try to examine all possible functional explanations concerning scent marking, but rather to test the generally accepted hypotheses (Gosling, 1982Gosling, , 1990 Gosling & McKay, 1990; Gosling & Roberts, 2001a, b) in regard to gerbils. ...
Article
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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.
... shelter and mates, are exclusive and actively defended by residents against conspecific intruders (Maher and Lott 1995). Territory defence typically involves the advertisement and maintenance of territorial boundaries through a suite of behaviours that may include scent marking, vocalizations and vigilance, and ultimately, chasing and fighting intruders (e.g., Krebs et al. 1978; Kacelnik et al. 1981; Gosling 1982; Packer et al. 1990; Lazaro-Perea 2001). These behaviours are expected to be costly, as they involve time, energy and, in some cases, risk to self (Davies 1980; Low 2006; Grinnell et al. 1995). ...
... Communication by way of scent marks (i.e., urine, faeces and scent gland secretions), may allow individuals to convey information regarding their sex, identity, group membership, health and reproductive status (Johnson 1973; Wyatt 2003). In the context of territorial behaviours, scent marking is expected to allow residents to advertise territory ownership and their competitive abilities (Gosling 1982; Wyatt 2003; Hurst and Beynon 2004). Scent marking may also be used by intruders as a form of advertising their presence to potential mates and as a challenge to territory owners (Gosling and Roberts 2001; Hurst and Beynon 2004). ...
... In the context of territorial interactions, scent marking is expected to allow residents to advertise territory ownership and their competitive abilities (Gosling 1982In Chapter 6, I showed that meerkats are able to discriminate between scent marks of extra-group males and scent marks from males within their own groups. All residents were more likely to emit alarm calls when investigating the scent of an extra-group male than the scent of a resident male. ...
Thesis
In group living animals where natal dispersal is delayed, prospecting allows individuals to asses their future dispersal and breeding opportunities and, in males of some species, may minimize the costs of delaying dispersal by enabling extra-group breeding while still resident in the natal group. While evidence of prospecting is widespread, comparatively little is known about the development of this behaviour and few studies have investigated the factors that may affect investment in prospecting, as it is typically difficult to monitor such mobile individuals. Prospectors typically encounter neighbouring groups during extraterritorial forays and resident individuals in these groups respond aggressively to approaches by extra- group males, given the potential loss in direct and indirect fitness that prospectors may inflict. As with prospecting behaviour, few studies have investigated the causes of individual differences in investment in repelling prospectors and measured the costs of such territory defence. In this dissertation, I exploit our ability to closely monitor prospecting males in meerkats, to investigate the causes of individual variation in extraterritorial prospecting effort and aggressive responses to prospector intrusions. In Chapter 3, I show that, as adults, heavier males invest more in prospecting than lighter ones, and that males time their forays in order to maximize their chances of dispersal, while minimizing the associated costs by prospecting when neighbouring groups are in close proximity to their own. In Chapter 4, I demonstrate that males that are heavier in early life start prospecting at a younger age and contribute less to helping later in life, than lighter males. In Chapter 5, I show that the threats posed by prospectors towards residents are associated with high investment by resident males in repelling intruders, which has measurable costs in terms of weight gain and cooperative contributions to offspring care. Finally, in Chapter 6, the experimental presentation of scent cues reveals that meerkats discriminate between resident and extra-group male scent cues, and that resident dominant males exhibit stronger responses to indirect evidence of prospectors than other group members.
... Scent-marking, the spatiotemporal pattern of scent deposition by an individual within a territory, is an important source of information. Scents deposited along trails, runways and prominent objects in the habitat not only allow the orientation of owners within their territories (Gosling, 1982) but also provide information to conspecifics about the competitive abilities of the owner (Fisher, Swaisgood, & Fitch-Snyder, 2003;. In territorial species, scent-marking allows to identify neighbours and consequently minimize aggressiveness during encounters ("dear enemy" hypothesis; Temeles, 1994). ...
... Only individuals that successfully defend a territory, and prevent intruders to overlay their own scents (Rich & Hurst, 1998), can ensure that their own scent predominates in it (Hurst, 1993). This also allows females to identify a male as the territory owner, by contrasting their body scents with the predominating scents in the area (Gosling, 1982(Gosling, , 1990Hurst, Thom, Nevison, Humphries, & Beynon, 2005;Rich & Hurst, 1998), and consequently assess the quality of a potential mate . In this way, chemical records of competitive interactions left on the substrate by scent-marking of a territorial male and by challenging countermarks of competitors are available to females which gain valuable information to discriminate between partners of varying quality. ...
... In the wild, competitive interactions would occur on a regular basis and individuals of higher competitive ability would be able to deter and countermark intrusions of low quality males. This information can be used by competitors to decide whether to challenge the owner or to flee (Gosling, 1982). But also, it can be used by females to evaluate the suitability of a male, as the Competitive Countermark Hypothesis (CCH) highlights . ...
Article
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.
... Based on a large number of studies, carried out on mammals and other terrestrial vertebrates, scent is known to play a major part in territorial defence, where it serves an extended phenotype role and is usually associated with paths and boundaries 6 . In addition, Gosling and co- workers 7,8 showed that territory holding mammals, including a number of deer and their relatives, also scent-mark their own bodies -the scent-matching hypothesis -which is thought to enable intruders to recognise territory holders and thus avoid fights they are likely to lose (while also protecting territory holders from the risk of damage). In previous articles, 9-12 we showed how fallow bucks concentrate their scent marking activities during the rut at conspicuous large scrapes located at the margins of mating territories. ...
... This challenger 'broad side' behaviour pattern appears to deviate from the classical Gosling model referred to above, which is based on the belief that body marking is restricted to territory- holders. 7 Moreover, the visually striking large buck display posture, which comprises a combination of splayed leg urination and overhead branch rubbing (Fig. 1D), was recorded during only 19% of total visits (N = 168), which is at variance with some early reports which gave the impression that this is the main form of scent marking behaviour employed by fallow bucks. Wallowing, previously thought not to occur in this species, 5 was of longer duration in the first half of the rut, which reflected the more stable, less stressful, conditions when the original territory holder was fully in command (first half: median length 19 min, range 1-190 min, n=17; second half: median length 1 min, range 1-11 min, n=16). ...
Article
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Scent-marking behaviour of male fallow deer (Dama dama), focusing on the role of the products of the preputial glands and urine, was investigated using animals inhabiting a private nature reserve in South Devon U.K. This publication deals with the olfactory behaviour of territory holders and non-territory holders, including the youngest cohort which appeared to use the scent of dominant bucks, derived from ground scrapes, for their own hierarchy-determining purposes, thus representing a possible example of cheating behaviour. The pattern of scent-marking recorded in this study did not fit with that expected from the classical scent-matching hypothesis.
... Based on a large number of studies, carried out on mammals and other terrestrial vertebrates, scent is known to play a major part in territorial defence, where it serves an extended phenotype role and is usually associated with paths and boundaries 6 . In addition, Gosling and co- workers 7,8 showed that territory holding mammals, including a number of deer and their relatives, also scent-mark their own bodies -the scent-matching hypothesis -which is thought to enable intruders to recognise territory holders and thus avoid fights they are likely to lose (while also protecting territory holders from the risk of damage). In previous articles, 9-12 we showed how fallow bucks concentrate their scent marking activities during the rut at conspicuous large scrapes located at the margins of mating territories. ...
... This challenger 'broad side' behaviour pattern appears to deviate from the classical Gosling model referred to above, which is based on the belief that body marking is restricted to territory- holders. 7 Moreover, the visually striking large buck display posture, which comprises a combination of splayed leg urination and overhead branch rubbing (Fig. 1D), was recorded during only 19% of total visits (N = 168), which is at variance with some early reports which gave the impression that this is the main form of scent marking behaviour employed by fallow bucks. Wallowing, previously thought not to occur in this species, 5 was of longer duration in the first half of the rut, which reflected the more stable, less stressful, conditions when the original territory holder was fully in command (first half: median length 19 min, range 1-190 min, n=17; second half: median length 1 min, range 1-11 min, n=16). ...
Article
Full-text available
Scent-marking behaviour of male fallow deer (Dama dama) during the rut, focusing on the role of the products of the preputial glands and urine, was investigated using animals inhabiting a private nature reserve in South Devon U.K. This publication deals with the olfactory behaviour of territory holders and non-territory holders, including the youngest cohort which appeared to use the scent of dominant bucks, derived from ground scrapes, for their own hierarchy-determining purposes, thus representing a possible example of cheating behaviour. The pattern of scent-marking behaviour recorded in this study did not fit with that expected from the classical scent-matching hypothesis.
... Sustained inspection of an unfamiliar outgroup odour may reflect a higher interest in a novel odour or increased efforts to obtain and process new information about the individual scent donor [68]. According to the scent-matching hypothesis, individuals of territorial species learn the scent of marks they encounter in the environment and then compare it with the scent of animals they meet, to facilitate appropriate behaviour [74,75]. Besides obtaining information about the state or identity of an intruder, chimpanzees might also be able to assess the distance to competing groups through the freshness or intensity of the odour or the frequency of outgroup scents encountered [74]. ...
... According to the scent-matching hypothesis, individuals of territorial species learn the scent of marks they encounter in the environment and then compare it with the scent of animals they meet, to facilitate appropriate behaviour [74,75]. Besides obtaining information about the state or identity of an intruder, chimpanzees might also be able to assess the distance to competing groups through the freshness or intensity of the odour or the frequency of outgroup scents encountered [74]. Although chimpanzees are not known to actively scent mark their territory, they sniff and inspect olfactory cues such as urine, faeces or traces of body odour in chimpanzee nests on boundary patrols [44,45]. ...
Article
Primates were traditionally thought to have a reduced sense of smell. Although there is now evidence that olfaction plays a greater role in primate social life than previously assumed, research on the sense of smell in non-human apes is scarce. Chimpanzees sniff the ground and vegetation on boundary patrols, but the function of this behaviour is unclear. Since chimpanzees are highly territorial and can kill individuals that do not belong to their own community, sniffing might function to gather information about conspecifics, particularly concerning group membership and kinship. To investigate whether chimpanzees recognize group members and kin via olfactory cues, we conducted behavioural bioassays on two groups of chimpanzees at Leipzig Zoo. In a pilot study, we found that chimpanzees responded more strongly to urine than to faeces or body odour. We then presented urine from group members, outgroup individuals and an unscented control in aerated boxes using a simultaneous discrimination task. The first behaviour after a chimpanzee first approached a box was related to olfaction (sniffing, nose within 20 cm, licking) in 83% of cases, highlighting the importance of olfaction as a general investigation mechanism in this species. Chimpanzees sniffed significantly longer at urine stimuli than the control and significantly longer at odours from outgroup individuals than those from group members. Furthermore, the duration of sniffing was positively correlated with relatedness. Our results suggest that chimpanzees use olfactory cues to obtain information about social relationships and fill a gap in our understanding of primate chemical communication.
... Although scent-marking and overmarking has been intensively studied in some mammalian taxa, especially rodents (Gosling 1982;Brown and McDonald 1985;Ferkin and Pierce 2007;Roberts 2007), other mammalian taxa remain understudied. Equids represent a very good group for studying this phenomenon for several reasons. ...
Article
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Overmarking occurs when one individual places its scent mark directly on top of the scent mark of another individual. Although it is almost ubiquitous among terrestrial mammals, we know little about the function of overmarking. In addition, almost all studies on mammalian overmarking behaviour dealt with adult individuals. Reports on this behaviour in juveniles are extremely rare, yet may elucidate the function of this behaviour. We tested four mutually non-exclusive hypotheses which might explain this behaviour in juveniles: (1) conceal the individual’s scent identity, (2) announcement of association with other group members, especially the mother—i.e., sharing identity with the mother, (3) to prevent the next conception of the mother, i.e., parent-offspring conflict, and (4) an early expression of male sexual behaviour. We observed 43 foals (out of 108 individuals) from all African equid species (Equus africanus, E. grevyi, E. quagga, E. zebra) in five zoos. In total, we recorded 3340 eliminations; 260 of these events were overmarked by 38 individual foals representing all species. This represents one of the highest rates of overmarking ever recorded by mammalian juveniles. Foals of all species except African wild ass overmarked the mother more often than another herdmate: with male foals overmarked at a higher rate than female foals. Mothers preferred to overmark foals, but not exclusively their own foal. Our results provide support for the hypotheses that overmarking serves to share identity between foal and mother, and that it is an early expression of male sexual behaviour.
... For instance, male snow voles (Chionomys nivalis) and house mice (Mus musculus) show less aggression towards males that have been recognised by " scent matching " than those that have not been recognised in this way (Gosling and McKay, 1990; Luque-Larena et al., 2001). Furthermore, because highly competitive individuals will typically produce more scent marks and mark out larger territories using olfactory signals (Gosling, 1982; Hurst and Rich, 1999), this type of scent matching behaviour can also be used to assess the likely outcome of a future encounter based on the spatial E-mail address: benjamin.charlton@ucd.ie and temporal pattern of scent marks (Gosling and Roberts, 2001). A prerequisite for scent matching to occur though is the ability to discriminate between the odour cues of different individuals. ...
... This distinction is important because it highlights the possibility that the ultimate goal of urine-washing might be to alter some physical or somatosensory property of the hands and feet rather than to deposit a scent (Harcourt, 1981 ). Some mammals, including many deer and some antelopes, also practice 'self-anointing' with urine, although in these cases the urine is typically sprayed onto the pelage or specialized scent glands rather than onto bare skin (reviewed by Gosling, 1982). Among primates, urine-washing has a curious taxonomic distribution that suggests its occurrence is determined in part by phylogeny. ...
Article
Some primates regularly wash their hands and feet with urine. This behaviour is outwardly similar to scent marking in other mammals, but it differs in that the urine is applied to the bare skin of the hands and feet rather than rubbed into the fur or applied directly onto an object in the environment. Empirical evidence for the functional significance of urine-washing remains inconsistent. We used rigorous statistical methods to examine environmental and social influences on urine-washing behaviour, using 4380 observation hours on five groups of wild white-faced ca-puchins (Cebus capucinus) in Costa Rica. Urine-washing frequencies were most strongly affected by environmental dryness, both within and between seasons, with markedly less urine-washing during humid conditions. Increased individual activity levels also promoted urine-washing. Among females, urine-washing was less frequent during lactation than during pregnancy and other reproductive states. Among males, urine-washing frequencies were greater in alpha males, who also exhibited a 'vigorous' form of urine-washing that may be functionally distinct. During the dry season, 3/5 groups exhibited more urine-washing than expected near fruit trees, but across groups there were no consistent spatial patterns for urine-washing with respect to water resources, home range overlap zones, core areas, inter-group encounter zones, and the home-range periphery. Urine-washing appears to differ fundamentally from common forms of mammalian scent marking. We suggest that its function is primarily mechanical, perhaps to apply a sticky residue to the hands and feet to improve grip on dry, arboreal substrates. Lesser signalling functions may include sexual signalling and resource labelling.
... A second hypothesis states that they provide a system of territorial demarcation, whereby faecal/urine deposits are placed around home range boundaries to act as a delineation of the territory, i.e. intergroup spacing (Brashares & Arcese, 1999;Stewart, MacDonald, Newman, & Cheeseman, 2001), similar to glandular scentmarking strategies observed in some lemur genera (Mertl-Millhollen, 1979. Although many mammalian species use latrines with the functional role of providing territorial defence, the deposition of glandular secretions in addition to faeces/urine can provide additional chemical information (Brown & MacDonald, 1985;Gorman, 1990;Gorman & Trowbridge, 1989;Gosling, 1982;MacDonald, 1980). As such, a third hypothesis is that latrines may strategically serve to advertise sexual condition via olfactory cues (Gorman, 1990;Woodroffe & Lawton, 1990). ...
Article
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Latrine behaviour is the repeated use of specific sites for defecation/urination, and is common among most mammals, including carnivores, herbivores, marsupials and rodents. While rare among primates, latrine use has been observed among some lemurs. It has been hypothesized that group-living primates may use latrines to maintain intergroup spacing (i.e. territorial defence) and for female advertisement of sexual condition. To test these, we conducted focal follows of three neighbouring southern bamboo lemur, Hapalemur meridionalis, groups in Mandena littoral forest of southeast Madagascar. From January to December 2013, we recorded all occurrences of latrine behaviour and characterized latrine sites to determine what factors influenced returning to specific latrines. Additionally, we attempted to elucidate the functional role of scent marking at latrines. We assessed the degree of home range overlap between neighbouring groups, and recorded intergroup aggression. Overall, latrines were almost exclusively visually conspicuous sites and located in the core areas of group home ranges. Best-fit models indicated that multiply visited latrines occurred more often in core areas, and were influenced by both sexes. Glandular scent marking at latrine sites was driven by males, and occurred more during the nonmating season. Males overmarked female scent-marks less often during the mating season and more often when younger males were likely to disperse. Thus, overmarking at latrine sites may function as a mate-guarding strategy to deter new males. Latrine use supports the energy frugality hypothesis, which proposes that lemur social systems, known for female social dominance and low rates of agonism, evolved as responses to the low productivity of Malagasy forests. The deposition of olfactory cues (i.e. faeces, urine, glandular secretions) at visually conspicuous sites may convey information to neighbouring conspecifics, thus reducing the need for intergroup agonism. Overall, latrine behaviour acts as a multi-modal means of intergroup communication.
... Conversely, the use of traditionally-rubbed trees by bears for intra-specific communication does not exclude the additional benefits of alleviating external irritants and offering points of orientation within a bear's home range. In short, our results are consistent with previous conclusions that rubbing is a form of social communication (Tschanz et al. 1970) that serves as a form of marking similar in function to that of other mammalian species (Gosling 1982). If tree rubbing is a form of marking behaviour, then the disturbance or removal of traditionally rubbed trees could disrupt communication among bears. ...
Article
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Tree rubbing or marking by bears has been observed throughout the northern hemisphere. Even so, this behaviour has rarely been studied. We documented 93 sites where grizzly bears Ursus arctos horribilis rubbed on 116 trees during 1986-1992, in the Yellowstone Ecosystem. We used logistic regression and information-based estimation and selection criteria to specify models that explained selection of sites and individual trees for rubbing by bears in our study area. The probability of rubbing peaked during May and June, the period of mating and moult, and declined thereafter. At the landscape level, grizzly bears selected for gentle south-facing slopes, forest/non-forest ecotones with sparse deadfall, and forest stands dominated by lodgepole pine Pinus contorta or Douglas-fir Pseudotsuga menziesii. Among the trees at sites where bears rubbed, we found strong selection for large diameters but no indication of selection for species. Rubbed trees were highly associated with travel routes likely used by bears, including game trails, recreation trails and forest edges. Rubbing was often oriented towards these likely travel routes. Short trails of entrenched pad-shaped marks leading up to rubbed trees were recorded at 58% of the sites where rubbing occurred. Contrary to reports of black bears Ursus americanus clawing and biting trees, we found shredded or bitten bark at only 9% of sites with rubbed or otherwise marked trees. Circumstantial evidence suggests that bears used trees primarily for rubbing their back and shoulders. Our findings are consistent with previous arguments that rubbing serves as a means of chemical communication.
... Communication by means of olfactory signals, particularly by scent marking, is a major mode of information transfer in mammals (for review see Brown & MacDonald, 1985; Eisenberg & Kleiman, 1972). It may function in territory or resource advertisement and defence, advertisement of social status and reproductive condition, social regulation, and mate attraction (e.g., Gosling, 1982; Gosling & Wright, 1994; Heise & Rosenfeld, 1999; Roberts & Dunbar, 2000; Rosell, 2002). Therefore, in more general terms, olfactory signals play a role in reproductive competition and mate choice and thus can be expected to be subject to sexual selection (e.g., Johnstone et al., 1997; Penn, 2002). ...
Article
Social communication and social monogamy Communication is an integral component of social processes and social evolution (Hahn&Simmel, 1974; Philips&Austad, 1990), and variation in social systems influences patterns of communication (Marler&Mitani, 1988). Amongst other functions, communication may serve to attract, stimulate, defend, or compete for mates, and thus is intimately related to social organization and mating systems (Hahn&Simmel, 1974; Johnstone, 1997). The intensity of signals and displays can serve as a cue to the quality of mates or opponents, and if interindividual variation of signal quality exists, sexual selection can act upon signals and displays through mate choice and intrasexual competition (Johnstone, 1995). Examples are visual and vocal displays in many mammals, birds, and other vertebrates (e.g., Clutton-Brock&Albon, 1979; Ryan, 1983; Hill, 1990). Since in most animals it is usually males who compete for females, and females who are choosy about males, sexual selection results in signals and displays being either exclusive to or exaggerated in males (Andersson, 1994). Nevertheless, mutual sexual selection can lead to signals being expressed to the same degree in both sexes (Jones&Hunter, 1993). This may be the case in monogamous animals, which are often monomorphic, both physically and behaviourally, including their communication patterns (Kleiman, 1977). Deviations from the behavioural monomorphism of signals and displays in socially monogamous animals can be expected if the selective forces driving males and females into a monogamous system differ between the sexes.
... Competitor assessment strategies might also be greatly enhanced by discrimination of individual odors: an individual that recognizes the scent of a high status individual may benefit if it can subsequently match that odor to the odor of an individual encountered in the environment (cf. Gosling 1982). This ability allows the individual to avoid potentially injurious escalation with a superior competitor without engaging in more direct probing of the individual's fighting ability and/or motivation. ...
Chapter
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This chapter reviews the recent studies revealing the role of chemical communication in regulating many giant-panda interactions. The identity of marking individuals, their sex and reproductive condition, and their social rank in the community are examples of the message that a panda may transmit via scent. The data presented highlights the importance of chemical communication to giant pandas, and suggests useful applications of scent in the conservation and management of the species, in both the wild and in captivity. There is a clear role in the application of chemical communication for captive breeding programs. The chapter also notes that scent communication in the giant panda is more complex than the simple monitoring of scent marks deposited in the environment.
... In line with the territorial defence hypothesis, the scent marking rate increases on days when intergroup encounters occur (Lledo-Ferrer et al., 2011;Roberts, 2012) and the spatial distribution of scent marks maximizes the probability of detection by an intruder (Gorman & Mills, 1984;Gosling, 1982;Gosling & Roberts, 2001b;Lewis, 2006;Schilling, 1980). This hypothesis suggests that scent marking is related to the size of the home range (Gorman & Mills, 1984). ...
Article
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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.
... Spraints carry information about characteristics of individuals, including their age, sex, and reproductive status (Trowbridge 1983;Kean et al. 2011Kean et al. , 2017. Additionally, marking behavior seems to be associated with resource defense (Gosling 1982;Kruuk 1992;Remonti et al. 2011). Otters deposit spraints on noticeable places, such as boulders, logs, or prominent tussocks of grass (Erlinge 1967;Eisenberg and Kleiman 1972;Kruuk 2006). ...
Article
Otters (Lutra lutra) mark territories by depositing feces on prominent spots, like rocks or large tussocks of grass. However, sometimes they defecate on intentionally mounded heaps made of mud and/or plant material. We show that in sites occupied by otters the frequency of heap-making significantly increases with the frequency of otters occurrence and is independent of the availability of naturally occurring potential marking spots. Constructing a heap is presumably costly, and thus heap-making could be an honest signal of propensity for defending territory. We propose that it should positively correlate with the quality of the territory and its owner.
... Males are more attracted to odours of oestrous females than of non-oestrous females ( Brown, 1979;Gorman and Trowbridge, 1989). As observed in other mammals, Civets tend to defend their territories for the purpose of reproduction and when the resources are in short supply ( Gosling, 1982;Richardson, 1993). The distribution of scent marked objects was high around the civetry sites. ...
Article
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Civet musk is one of the natural resources in Ethiopia. Much as Ethiopia is the world's largest supplier of civet musk to perfumery industry in Europe, its Civet farming practices and knowledge about Scent-marking patterns of African Civet needs to be regulated for acceptable welfare standards and important to increase musk production. The scent marking objects and its preference were known by making 70 quadrates of 20 X 10 m 2 which were laid in different parts of the study area. This was done based on the proximity of the area to the latrine site, In the present study area, the African Civets used plants, metallic objects and poles to scent mark. Out of the 92 scent marking sites observed, 96.72% was within 100 m radius of the civetry. About 75% of the scent marks was observed within < 4 m distance from the path way. There was a high level of preference to mark on Bedena tree (Balanites aegyptiaca) (38%) followed by metallic objects (19.56%). In the wild, the amount of perineal gland secretion collected from each marked site varied from 0.0047 g to 0.98 g.
... Whatever the mechanism, successful recognition requires the existence of reliable indicators of the potential threat an individual represents. In mammals, scent-marking provides information about territory occupancy and serves as a preventative measure for territory defense (Ralls 1971;Gosling 1982). ...
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Territorial animals are expected to adjust their response to intruders according to the perceived threat-level. One of the factors that drives threat-level is the identity of the intruder. The dear enemy phenomenon theory postulates that individuals should respond with lower intensity to neighbors, already possessing a territory, than to strangers that may fight to evict them. In social species, the hierarchical status of the intruder might also mediate this response. Such behavioral adjustments presuppose a capacity to discriminate between individuals posing different threat levels. Here, we tested the behavioral response of Alpine marmots to territorial intrusions in a wild population. We compared both dominant females and males responses to scents from neighbor and stranger dominant males (dear enemy phenomenon) and to dominant and subordinate stranger males (social status-specific response). In addition, we tested for any covariance between male scents and social status. We showed that female and male dominant marmots do not adjust the intensity of their behavioral responses to whether the intruder's territory is bordering or not (neighbors or strangers) or to the intruder's social status, even though dominant and subordinate males are thought to pose different threats and social status is encoded in scents. Thus, we did not find support for the dear enemy phenomenon and conclude instead that, in dominant Alpine marmots, no intruder should enter a foreign territory. Research taking a more holistic approach of the evolution and maintenance of territoriality is required to understand the flexibility of responses to intruders in group-living species.
... Unlike acoustic and chemical signals, which may reach a potential prey or a predator, even if its view is obstructed [18,19,20], visual signals decrease the possibility of prey/predator intercepting a message intended for closer conspecifics warning about their presence. In turn, acoustic and chemical communication play important roles in advertising territory occupation and ownership [18,20,21]. This highlights white marks, as well as acoustic and chemical communication, as functional traits that may increase the fitness of organisms and/or their influence on other organisms and on ecosystem functions [22]. ...
Article
Melanism in the cat family has been associated with functions including camouflage, ther-moregulation and parasite resistance. Here we investigate a new hypothesis proposing that the evolution of melanism in cats has additionally been influenced by communication functions of body markings. To evaluate this hypothesis, we assembled a species-level data set of morphological (body marks: white marks on the backs of ears) and ecological (circadian activity: arrhythmic/nocturnal, and environmental preference: open/closed) characteristics that could be associated with communication via body markings, and combined these data with a dated molecular phylogeny. Next, we tested the association between melanism and communication, first by relating species' body marks with their ecological conditions, using a Bayesian implementation of the threshold model. Second, to explore the evolution of characteristics potentially influencing melanism in cat species, we modeled their evolution relative to melanism using models of coordinated vs. independent character changes. Our results suggest that white marks are associated with intraspecific communication between individuals that have non-melanistic phenotypes, as well as towards melanistic individuals (without white marks). The absence of white marks in a melanistic individual tends to be a limiting condition for intraspecific visual communication at night, resulting in an evolutionary dilemma for these species, i.e. to be almost invisible at night, but not to communicate visually. The comparative analysis of several evolutionary models indicated more support for the evolution of melanism being coordinated with the evolution of arrhythmic activity and white marks on the backs of ears.
... In spite of our limited sample size and of captive conditions, a few preliminary conclusions may be drawn. Although intense marking was more evident in the male sex, females showed it too, confirming the prediction of territorial marking in both sexes [8,9]. Most marking occurred using the secretion of preorbital glands, as it has been reported for the Formosan serow C. swinhoei [42]. ...
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Comparative behavioural studies help reconstruct the phylogeny of closely related species. In that respect, the serows "Capricornis" spp. occupy an important position as they have been assumed to be the closest forms to the ancestors of Caprinae. In spite of that, information on the behavioural repertoire of the mainland serow "Capricornis sumatraensis" is exceedingly poor. In this paper, we report data on the activity rhythms and social behaviour of rutting mainland serows in captivity (Central Thailand, January 1986; January–February 1987). Activity was bimodal with peaks in mid-afternoon and late night. Resting and ruminating peaked at noon and twilight. Four patterns of marking behaviour were observed out of a total of 1900 events. Males and females were found to use di�erent marking sites and frequencies. A total of 33 social behaviour patterns were observed: 18 patterns concerned agonistic behaviour, whereas 15 patterns were relevant to courtship behaviour. A comparison across Caprinae species with unritualised piercing weapons (i.e., Capricornis, Naemorhedus, Rupicapra, Budorcas, and Hemitragus) has shown that inter-sexual direct forms of aggressive behaviour are used significantly more often than indirect ones, but for chamois, confirming "Rupicapra" spp. as the most advanced genus among them in terms of an early ritualisation of weapons. Conversely, horns of the goral "Naemorhedus" spp. and the serow lie on the same plane of the frontal bones, thus making possible the usage of a dominance display through frontal pushing.
... Numerous studies have shown that in territorial defence threats, song and scent marking have a number of advantages over violent combat, because of e.g. elimination of the risk of injury, and so death (Johnson, 1973;Brown, 1975;Maynard Smith, 1982;Gosling, 2010), lower energetic costs (Brown, 1975) or less reduced predator vigilance (Dunn et al., 2004). ...
Article
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... Some mammals use scent-marking to convey information about their reproductive and social status to other individuals in their population, as well as to mark their territories (Gosling, 1982;Rodgers et al., 2015;Allen, Gunther & Wilmers, 2017;Wachter et al., 2017;Wikenros et al., 2017). This form of communication is reliable as the information remains long after the individual has left, making it ideal for solitary carnivores (Cornhill & Kerley, in press). ...
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Competition occurs between species for shared resources. Subordinate species employ resource selection to shift their resource use away from that of dominant species in order to avoid the negative consequences of competition. Only in Africa is the post-Pleistocene large carnivore guild intact, consisting of lions Panthera leo, spotted hyaenas Crocuta crocuta, leopards Panthera pardus, cheetahs Acinonyx jubatus, and African wild dogs Lycaon pictus. Therefore, only in Africa can we explore how large carnivores co-exist with one another in a diverse community. Cheetahs are a subordinate member of the large carnivore guild due to their small stature and solitary nature. However, we still do not fully understand how competition shapes cheetah behaviour and resource use. I used cheetahs as a model subordinate predator to determine the behavioural responses and resource selection of cheetah in response to assumed competition from other large carnivores. I experimentally explored the behavioural response of cheetah to large carnivore scent sources (scats) and their presence at cheetah scent-marking sites to test for avoidance of such cues. Moreover, using unplanned experiments based on the resource use of cheetahs in the absence and then presence of lions (assessing space and time use by cheetah) and African wild dogs (assessing prey use by both species), I evaluated resource selection by cheetahs as a way to reduce competition with these large carnivores. In addition, I measured spatial and temporal partitioning between cheetahs and all four large carnivores at camera trap sites. Finally, I assessed whether cheetahs responded to competitors using a long term proactive response or a short term reactive response. I found that all four of the other large carnivores in the guild shaped cheetahs resource use, however, all did so on different axes in accordance with the niche complementarity hypothesis. Lions and spotted hyenas were avoided through time, African wild dogs through space and prey use, and lions and leopards on the spatiotemporal axis. Moreover, I show that cheetahs utilize a reactive response to competition that allows them to avoid risk while still obtaining necessary resources.
... Based on our results (Fig. 1), dominant hamsters exhibited more chasing behavior, while subordinate hamsters exhibited more fleeing behavior after a 1-week intense conflict. Scent rubbing was previously shown to be associated with social status advertisement, which can be assessed by subordinate individuals to reduce agonistic costs [52]. Hormones released by HPA system are able to interact with gut microbiota in colon under stress conditions [53], while corticosteroid will decline after a dramatic increase during the initial stage [54]. ...
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Social stress can dramatically influence the health of animals via communication between gut microbiota and the HPA system. However, this effect has been rarely investigated among different social ranked animals after chronic repeated social encounters. In this study, we evaluated changes and differences in microbiota among control, dominant, and subordinate male greater long-tailed hamsters (Tscherskia triton) over 28 successive days of repeated social encounter. Our results indicated that as compared with the control group, short-term repeated social encounters significantly altered fecal microbiota of subordinate hamsters, while chronic repeated social encounters altered colonic mucosa-associated microbiota of both dominant and subordinate hamsters. Fecal microbiota showed a transition in composition and diversity on day 2 for the subordinate group but on day 4 for the control and dominant groups under repeated encounters. Compared with their baseline, genus Lactobacillus increased in both dominant and subordinate groups, while genus Bifidobacterium increased in the subordinate group and genus Adlercreutzia increased in the dominant group. Our results suggest that chronic repeated social encounter can alter diversity and composition of gut microbiota of hamsters in both feces and colonic mucosa, but the latter performed better in reflecting the effects of chronic stress on microbiota in this species. Future studies should focus on elucidating how these microbiota alterations may affect animal behavior and fitness.
... Unlike acoustic and chemical signals, which may reach a potential prey or a predator, even if its view is obstructed [18,19,20], visual signals decrease the possibility of prey/predator intercepting a message intended for closer conspecifics warning about their presence. In turn, acoustic and chemical communication play important roles in advertising territory occupation and ownership [18,20,21]. This highlights white marks, as well as acoustic and chemical communication, as functional traits that may increase the fitness of organisms and/or their influence on other organisms and on ecosystem functions [22]. ...
Article
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Melanism in the cat family has been associated with functions including camouflage, ther-moregulation and parasite resistance. Here we investigate a new hypothesis proposing that the evolution of melanism in cats has additionally been influenced by communication functions of body markings. To evaluate this hypothesis, we assembled a species-level data set of morphological (body marks: white marks on the backs of ears) and ecological (circadian activity: arrhythmic/nocturnal, and environmental preference: open/closed) characteristics that could be associated with communication via body markings, and combined these data with a dated molecular phylogeny. Next, we tested the association between melanism and communication, first by relating species' body marks with their ecological conditions, using a Bayesian implementation of the threshold model. Second, to explore the evolution of characteristics potentially influencing melanism in cat species, we modeled their evolution relative to melanism using models of coordinated vs. independent character changes. Our results suggest that white marks are associated with intraspecific communication between individuals that have non-melanistic phenotypes, as well as towards melanistic individuals (without white marks). The absence of white marks in a melanistic individual tends to be a limiting condition for intraspecific visual communication at night, resulting in an evolutionary dilemma for these species, i.e. to be almost invisible at night, but not to communicate visually. The comparative analysis of several evolutionary models indicated more support for the evolution of melanism being coordinated with the evolution of arrhythmic activity and white marks on the backs of ears.
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Animals exploring a new environment develop cognitive maps using diverse sensory input and, thereby, gain information needed to establish home ranges. Experiencing, and learning information about, resources should be advantageous to the resident of a home range while lack of such information should put invaders into the home range at a disadvantage. Conspecifics, especially, should avoid the home ranges of one another to ensure that they do not experience reduced resource availability caused by resource depression or depletion. Yet, encountering conspecific competitors of different sexes may elicit responses that can lead to spacing on a landscape that has different costs and benefits on males and females. We tested the hypothesis that female fishers ( Pekania pennanti ) avoid competition from both males and female conspecifics whereas male fishers avoid competition only from other males. We reintroduced fishers onto our study site in the presence or absence of competitors’ home ranges during late 2009 through 2011. Using satellite transmitters (Argos) and land-based (VHF) telemetry, we monitored fishers and estimated their locations, movements and use of the surrounding landscape during their first 500 days after release. All fishers settled in relatively high-quality habitat but females that encountered the home ranges of conspecifics moved farther, explored larger areas, and settled farther from their release locations than did females that did not encounter a conspecific’s home range. Male fishers exhibited diverse responses upon encountering the home ranges of conspecifics. Thus, female fishers avoid conspecific competition from all fishers, but males tolerate, or impose, competition with females, apparently to increase mating opportunities. These observations are consistent with the movements and strategies of other solitary carnivores.
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Ecological psychology is built on a perception-oriented ontology. The primary focus has been on explaining the perception and action behavior of individual animals. To accommodate social phenomena within the ecological approach, it is necessary to expand the ontology, however theorists have been unclear about how to do this. The paper presents a negative argument and a positive programmatic outline. The negative argument is against the use of the term ‘social affordance’, a term that confuses the perspective of the researcher with that of the animal. Instead, it is advocated that we adopt, as a working hypothesis, the claim that all affordances are social; that is, all affordances are public and are, in principle, observable by a third party. The programmatic outline then shows that affordances alone are insufficient for describing social meaning. An ecological social ontology requires new tools for describing interaction processes, symbolic meaning, and material culture as structures occurring within the populated environment.
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Much of the growing interest in avian chemical signals has focused on the role of kin recognition or mate attraction, often with an emphasis on males, with uropygial gland secretions perhaps providing information about an individual’s identity and quality. Yet, data collected to date suggest sexual dimorphism in uropygial glands and secretions are often emphasized in female, rather than in male birds. That is, when a sexual difference occurs (often during the breeding season only), it is the female that typically exhibits one of three patterns: (1) a larger uropygial gland, (2) a greater abundance of volatile or semi-volatile preen oil compounds and/or (3) greater diversity of preen oil compounds or associated microbes. These patterns fit a majority of birds studied to date (23 of 30 chemically dimorphic species exhibit a female emphasis). Multiple species that do not fit are confounded by a lack of data for seasonal effects or proper quantitative measures of chemical compounds. We propose several social functions for these secretions in female-based patterns, similar to those reported in mammals, but which are largely unstudied in birds. These include: (1) intersexual advertisement of female receptivity or quality, including priming effects on male physiology, (2) intrasexual competition, including scent marking and reproductive suppression or (3) parental behaviors, such as parent-offspring recognition and chemical protection of eggs and nestlings. Revisiting the gaps of chemical studies to quantify the existence of female social chemosignals and any fitness benefit(s) during breeding are potentially fruitful but overlooked areas of future research.
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The intensity of scent marking by territorial beavers could relate to different numbers of transient beavers. Rivers are, most likely, the main paths of beaver dispersal. It is supposed that the intensity of subadult migration depends upon the size of the river because those basins of larger rivers potentially contain a greater number of beavers. Scent marking intensity (the number of scent mounds) and relative abundance (number of beaver activity signs) of beavers were studied in 18 segments of various rivers in Lithuania during the period of subadult dispersal during April and the first ten days of May. The total length of transect was 157 km. The yield of water (Q) of these river segments varied from 0.4 to 32.2 m3/s. The Lithuanian beaver population is considered abundant. Even the smallest peripheral water bodies are densely inhabited by beavers, thus making the dispersal of population surplus rather complicated. The highest scent marking intensity (1.78 scent mounds per 0.25 km of shore line) was found in medium-sized rivers (mean Q = 4.6 m3/s), and the lowest (0.64 scent mounds per 0.25 km of shore line) in the smallest rivers (mean Q = 0.6 m3/s). Scent marking intensity was significantly positively correlated with the relative abundance in most rivers, excluding the smallest ones. The results are discussed in relation to an appropriate response of territory owners to the different intensity of subadult migration in rivers of differing sizes. The scent mound system seems to be an important mechanism of population self-regulation, preventing an overexploitation of ecological resources in rivers where intensive migration of beavers occurs.
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From an evolutionary viewpoint, signals generally should be reliable or honest (Zahavi, 1987; Johnstone, 1997). Animals can gain a number of advantages from advertising high competitive ability to potential mates and to other competitors, particularly males which often compete strongly for mating opportunities (Andersson, 1994). Animals often prefer high quality mates that will increase the fitness of their offspring, both because of genetic benefits (through good genes or Fisherian selection) and because parents of high competitive ability often provide better resources and protection. Competitors will also gain an advantage if potential challengers withdraw from, or otherwise avoid, aggressive encounters with an opponent of high fighting ability. There is thus strong selection pressure on signallers to advertise high social status and competitive ability to others. However, receivers will only gain an advantage from responding to such signals if these are reliable indicators of the signaller’s competitive ability. Females that mate with low quality males that dishonestly signal high competitive ability will gain no advantage for their offspring, while males that withdraw from agonistic encounters with poorer competitors will be disadvantaged. There is thus strong selection on receivers to respond only to honest signals that are resistant to cheating, and therefore for high quality animals to provide such reliable information in signals as these will be effective in attracting mates and deterring competitors.
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Females in promiscuous species (e.g. house mice; Mus musculus domesticus) may mate with multiple males, which can negatively influence the reproductive success of males because of the risk of sperm competition or injury due to male-male competition. Thus, selection should favor those that can discern not only if a female has recently been in contact with another male (potential risk of sperm competition), but whether the other male with which the female had been in contact is dominant (potential risk of injury) or subordinate to the investigating male.
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Communal marking sites, or latrines, appear to play an important role in intraspecific communication and social dynamics in a wide range of mammal species. The spatial distribution of latrines can provide clues to their function and has been well documented in a number of species. Latrine use may vary considerably through time, however, and a more comprehensive approach to their study that considers spatial and seasonal patterns of use is required to understand more fully the costs and benefits of latrine use, and hence their adaptive significance. This study investigated spatial and seasonal patterns of latrine use by spotted hyena (Crocuta crocuta) in northern Botswana, examining their potential role in resource defense. Latrine characteristics and hyena activity were monitored to test the influence of season and location (relative to clan territories and roads) on latrine use. We conducted monthly scat counts (at 78 latrines) and continuously recorded hyena visitation (to 50 latrines) in five clan home ranges, demonstrating clear seasonal patterns in latrine use. Latrines were smaller in the wet season (November-March), resulting from fewer visits by hyenas, reduced scat accumulation, and the seasonal activity of coprophagous beetles. We speculate that such a seasonal pattern may be driven by reduced competition for food during the wet season. Latrines located within core clan areas were no larger or more frequently used than those in home-range boundary areas, but hyenas did preferentially place latrines alongside roads and were more likely to reuse road-side latrines in subsequent years. This pattern was not due to observer detection bias and adds to the growing body of literature on the impact that roads and other anthropogenic features have on the communication and movement ecology of wild animals. Significance statement Although most mammals use communal marking sites, very little is known about their function and detailed patterns of use in many species. We investigated latrine use in spotted hyena (Crocuta crocuta) and describe spatial and temporal marking patterns that are consistent with optimizing scent longevity and detection. Spatially, hyenas preferentially located latrines along the edge of man-made vehicle tracks, which may increase signal detection and transmission. Seasonal marking patterns suggest that hyenas optimize their communication by concentrating activity in the dry season, thereby avoiding the disruptive effects of coprophagous dung beetles and rain, and focusing activity during periods when food resources are expected to be scarcer. These results demonstrate seasonal and spatial optimization of communication, including in response to novel anthropogenic features in the environment such as roads, advancing our understanding of communication strategies in mammals more broadly.
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In order to investigate and create an ethogram of the rolling behavior of mules (Equus caballus × Equus asinus) in comparison to that of horses (Equus caballus), we observed a domestic free-range, mixed group of 17 horses and 11 mules for two consecutive days (12 h/day), filming all the rolling events. From these footages, we were able to register the occurrence of specific behaviors that characterize the rolling: sniffing the ground prior to lie down; tail swishing; resting in recumbence; scratching and shaking. Moreover, we also observed two social dimensions that are possibly involved with the rolling behavior: 1) Social facilitation - once an animal started to roll, others in the group copied its behavior and 2) Demarcation - animals tended to return to roll in the same dust patches. In this equine group, mules rolled more frequently than horses and also exhibited more of the specific behaviors tail swishing, resting and scratching. Moreover, in the rolling events exhibited by the mules, the social facilitation and demarcation occurred more than in the horses. As a preliminary study, we pointed some important differences between the rolling behavior of horses and mules. Considering the lack of knowledge in this regards such findings could be useful to spread ideas for future studies concerning the mules behavior and to improve the welfare conditions of these animals.
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In mammals, a low molecular mass protein (17-20 KDa) reported from the pheromone sources such as urine, saliva, glandular secretion, etc., as ligand-carrier (pheromone carrier) has been associated with chemo-communication. Since the preorbital gland post is one of the major pheromone sources in Indian Blackbuck, an endangered species, we assumed thta it possibly contains low molecular mass protein for chemical communication. Hence, we investigated the preorbital gland post in territorial and non-territorial male blackbucks for such low molecular mass proteins adopting SDS-PAGE and LC-MS/MS analysis. The total content of protein was higher in the post of territorial males than non-territorial males of adult and sub-adult. In fact, the protein profiles such as 17, 21, 25, 42 and 61 kDa were noted in the gland secretion of territorial and non-territorial males. The intensity of the 17 kDa protein band was higher in territorial males than nonterritorial males. In-gel trypsin digestion of the 17 kDa band was processed and subjected to LC-MS/MS and SEQUEST analyses. The results of LC-MS/MS and SEQUEST search showed the presence of α2u-globulin in the 17 kDa band. In addition, the identified α2u-globulin sequence possessed GDW residues, which are the characteristic signature for lipocalin family. Since the α2u-globulin has been reported from the pheromone-carrying proteins in some mammals, this protein may carry the volatiles (pheromone compounds) in male Blackbucks preorbital gland to evoke the scent marking for maintaining territoriality (home range) and attraction towards female, through the secretion of glandular protein. © 2015, National Institute of Science Communication. All rights reserved.
Chapter
Human–wildlife conflict (HWC) is a pervasive, global conservation issue that occurs whenever interactions between people and wildlife result in a negative outcome for either or both parties or the resources upon which they rely. Humans are impacted by wildlife either directly through their survival or health, or indirectly through damage to their food resources or property. Alleviating HWC has relied upon means from the individual to the population scale. Problem animals can be killed, relocated, or trained to avoid human habitations. Animal populations can be reduced through lethal or non-lethal means. Lethal control or relocation often have consequences such as biodiversity loss, unforeseen trophic cascades, and other ecologically detrimental effects. In many cases, the preference may be to keep the animals in question in the region but to reduce their impact on humans. In such situations, barriers can be constructed but these also have potential negative ramifications for other species as well as high costs for construction and maintenance. For invertebrate pests, a push–pull integrated pest management strategy has become increasingly popular. Chemical signals in the form of a natural repellents and attractants serve to keep pests away from crops and attract them to other areas, often traps. For vertebrates, the lack of known chemical signals hampers this approach and furthermore, the lethal trapping of individuals often is undesired. However, biological fences in concert with training animals to avoid the conflict area (e.g., crop or livestock) have potential for reducing HCW, especially for longer-lived, medium- to large-sized vertebrates. In lieu of using natural signals with known function, the proposition is to invoke a novel odor that would be unusual in the range of the problem species. Such an odor would be associated with strongly aversive stimuli near points of conflict and used as a warning of such a stimulus for future encounters. This novel scent fence would elicit anxiety and arouse a memory of discomfort and fear in the animal, motivating it to move away from the area without another interaction with the aversive stimulus. The odor would create an atmosphere of unease and avoidance rather than be the repellent itself. This approach provides further impetus for understanding better the chemical communication and the specific chemical signals used by vertebrates. In advance of those discoveries, odors may be useful as part of an aversive conditioning program to alter movement patterns of animals and keep them away from crops and livestock.
Chapter
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The mating strategies of male mammals consist of intrasexual competition for access to females and then attempts to maximise contacts with receptive females. The form that these attempts take is strongly influenced by female movements in response to food and their tendency to form social groups. Female reproductive success (RS) is primarily (although not entirely) determined by the number of offspring that can be produced and the factors that limit this are the availability of the nutrients needed for reproduction and the chance that offspring will be killed by predators. Many aspects of female behaviour are profoundly influenced by these considerations: for example, the movements of female antelopes in relation to spatial and temporal variation in grassland quality appear to be adaptations to optimise food quality while avoiding habitats that offer cover to predators. In addition, females can reduce the chance of predation by cryptic behaviour, alone or in small groups, or by the “selfish herd” advantages of joining larger social groups. Collectively, these female behaviours appear to dictate the mating strategy that is most profitable for males to adopt (Gosling, in press). Excluding lekking, these strategies fall into two main classes: (1) males follow one or more females, waiting until they become receptive, and (2) they defend part of the food resource that females need so that they can intercept females that are attracted to the resource (Gosling, in press). This second strategy is known as “resource defence territoriality” (Emlen and Oring, 1977).
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Identifying specific signaling components within complex mixtures is a common problem in semiochemistry. Both glandular secretions and excretory products contain components of semiochemical importance, but identifying these signals is problematic because they are usually parts of mixtures with several 100 components, of which only a subset may be involved with signaling. In contrast to waste and metabolic byproducts—which can be expected to vary both between and within individuals according to extrinsic factors—signaling compounds are expected to be uniform among animals sending the same signal and stable over time. In group-living territorial species we would expect there to be a degree of group-specificity in signals that advertise territory residence. As part of an ongoing study investigating and manipulating scent-marking and territorial behavior in African wild dogs, several 100 volatile components have been located from their urine. How many and which, if any, of these have active roles in semiochemical communication of territory residence is currently unknown. Observations of scent marking behaviors of African wild dogs strongly suggest that dominant urine overmarks (DUOs)—where one member of a pair deposits urine on the urine of its partner—are the most likely source of such signals. We used multivariate statistics to investigate >990 separated chemical components (some of which could be multiple compounds) found in these DUOs, and found as few as 10 chemical components that together enabled statistical discrimination of specific dominant pairs. We suggest that this method may be broadly applied across communication systems to locate components of signals within complex “mixtures.”
Article
Signal detection theory predicts that animals should select scent‐marking sites in a way that maximizes their probability of detection by target receivers. Many studies have been conducted with a focus on signaling behavior and function. Yet, the role of the environment in structuring scent‐mark patterns is poorly known. We studied the giant panda, a solitary species that relies heavily on chemosignaling for communication through depositing urine and anogential gland secretions (AGS) on trees, to determine how environmental factors influence scent deposition strategies. Using a combination of sign surveys, camera trapping, and gas chromatography mass spectrometry analysis, we conducted a comparative study of giant panda scent marking in two reserves, Wanglang and Foping, in China. The two reserves differed with regard to prevalence of tree species that differed in bark roughness, diameter at breast height (DBH), and moss status. For urine‐marking pandas selected large, mossy trees with very rough bark trees, whereas AGS marking was used preferentially on rough‐barked moss‐free trees in both reserves. Both AGS and urine contained more volatile than nonvolatile constituents and urine had higher relative abundance of volatiles than AGS, indicating urine likely has a larger signal range and shorter signal persistence than AGS. Urine marking was significantly more prevalent in Wanglang, which contains more rough‐barked, moss covered, and higher DBH trees favored for urine marking, than in Foping. Thus, habitat features appear to constrain pandas’ use of different types of marking, potentially altering how the communication system functions in different ecological contexts. Our study underscores the importance of future study evaluating environmental influences on chemosignaling behavior. Signal detection theory predicts that animals should select scent marking sites in a way that maximizes their probability of detection by target receivers. Many studies have been conducted with a focus on signaling behaviour and function. Yet, the role of the environment in structuring scent mark patterns is poorly known. We conducted a comparative study of giant panda urine and anogential gland secretions scent‐marking in two reserves, Wanglang and Foping, in China, to determine how environmental factors influence scent deposition strategies. We found that urine marking was significantly more prevalent in Wanglang, which contains more rough‐barked, moss covered and higher diameter‐at‐breast‐height trees favored for urine marking, than in Foping. Habitat features appear to constrain pandas’ use of different types of marking, potentially altering how the communication system functions in different ecological contexts.
Article
Although several mammals impregnate their fur with environmental odors, a phenomenon termed scent anointing or rubbing, the functional relevance of this behavior often is unclear. One theory is that scent anointing could be a form of scent matching with environmental odors to signal competitiveness and home range occupation. In this study we presented giant pandas with a range of odors to determine whether scent matching could provide a functional explanation for scent anointing in this species. We found that only a musk-based perfume elicited significantly more scent-anointing and scent-marking behavior than control. Males were also significantly more likely to scent-anoint and scent-mark than females. A preference for anointing, but not scent marking, when presented with peppermint (an insecticide) also was revealed. Our results suggest that giant pandas differentially scent-anoint with foreign odors to signal home range occupation, and possibly to repel ectoparasites. We also highlight how chemical signaling of resource-holding potential is likely to play an important role in determining competitive interactions between adult male giant pandas.
Chapter
This chapter introduces main concepts of rodent olfactory neuroethology and draws upon some particularly elegant examples of studies in rodents within natural and semi-natural environments to explore principles of ethologically-relevant rodent olfactory communication. As an initial framework for the understanding of rodent olfactory communication, it is useful to consider that rodents emit chemical signals which may be received by either members of their own species, or members of other species. While the emission of olfactory signals may have initially evolved for intraspecific purposes, the release of these cues can have unwanted consequences. Alternatively, in a more complex sequence of events, perception of a predator odor signal or perception of an odor from a stressed conspecific may result in an "alarm" signal in rodents to aid conspecifics in survival. These forms of olfactory communication are referred to as interspecific. The chapter finally outlines examples of how rodents utilize olfactory communication methods.
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This paper deals with rutting behaviour in two groups of reindeer studied at the Reindeer Research Station in Swedish Lapland. The first visible signs of rutting behaviour are described as well as the establishment of the harem and its ruler. Old bulls copulate more often than younger ones owing to the earlier occurrence of rut in these, their stronger fighting ability and the fact that old bulls are more polygamous than their younger rivals.The chasing away of intruders, and the constant herding of his own herd, keeps the ruling bull very active. Since, furthermore, he does not eat he becomes very thin during the rutting season.The size of the area necessary for a harem varies. A big bull with a large harem is able to keep his rivals away from a considerably greater area than a young bull with a small number of cows.The rutting territory seems not to be defined by particular geographical boundaries, but is an area, which changes with the movement of the harem and has no topographical references. The author has observed nothing that indicates territory markings in the proper sense, but the bull urinates on his hind legs, especially at threatening and exciting moments, and this behaviour may be interpreted as a form of territory marking.The rutting sound of the bull is a series of husky, rapid, rather low-voiced rattles that are brought about by the air being repeatedly pressed out of the lungs. The sound differs slightly from that made when the bull is driving in the immediate vicinity of a cow.Threat behaviour has been described as well as overflow activities resulting from a high level of aggressive behaviour. A typical rutting behaviour, that could be characterized as courting, is the bull's driving of the cows, i.e. the bull's running or walking after a cow depending on her heat status.Aggressiveness on the part of the ruling bull is more evident against older rivals than against younger ones, and old cows are particularly aggressive against young bulls.The first copulations usually take place during the last days of September. Mating activity reaches it peak at dusk and at dawn. Copulation seems to take place only once with each cow. The heat development of the cow has been described as well as the copulation act.
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Dominant and subordinate Harris' sparrows usually coexist in the same flock and signal their social status by a distinctive plumage badge. We investigated advantages of subordination and costs of dominance in an effort to understand both why dominants and subordinates coexist when groups are not composed of relatives and why an inexpensive signal of high status should not be mimicked by poor fighters. Dominants and subordinates do not seem adapted to different microhabitats determined by seed density. Instead, we hypothesize that they coexist in a manner analogous to shepherds and sheep with dominants defending space in good habitats for subordinates and subordinates being used as food finders by dominants. In support of this shepherds hypothesis dominants displaced other dominants more frequently than they displaced subordinates when neither bird was feeding, but dominants displaced any lower ranked birds from caches of seed. This difference in aggression at and away from small and hidden patches of concentrated food suggests that a cost of dominance is fighting with other dominants and that a benefit of subordination is not having to fight with dominants for space in good habitats. Showing this helps explain why subordinates signal their subordination by a plumage badge. Curiously, among nonfeeding birds subordinates attack subordinates more frequently than do dominants. This indirect interference competition among subordinate Harris' sparrows could be for the privilege of being "sheep" of dominant Harris' sparrows or for the privilege of being "shepherds" to other lower ranked species of the community. This competition among subordinates (and, by extrapolation, within plumage or status ranks) helps explain why Harris' sparrows are fairly continuously variable in coloration rather than dichromatic with respect to dominance status.
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An attempt has been made to distinguish between action area, home range, and territory in the Bovidae. The establishing of subjective boundaries is considered to be the most important criterion of territoriality. The existence of such boundaries becomes evident from certain behavioural symptoms; “defence” or better, localized dominance which may lead to intolerance, is one of them.Not all bovids are territorial. Within the territorial species, there seem to be at least two types: (a) The animals, usually in pairs, may, under favourable conditions, stay in their territories permanently; (b) Only the males are territorial and stay in temporary territories, usually for several weeks or months. This last type is obviously more common in horned ungulates than the first one. Within this second type (b), there are species-specific differences. For example, in Grant's gazelle (Gazella granti), under certain environmental conditions, this type of territoriality is combined with harem behaviour, but in the co-inhabiting Thomson's gazelle (Gazella thomsoni), the females roam through the territories of the males and stay together with the same buck only for a few hours per day.Even within one and the same species, there can be variations, apparently linked with differences in environmental conditions. This is discussed, using the examples of the Uganda kob (Adenota kobj, the wildebeest (Connochaetes taurinus), and Grant's gazelle. Finally, there can be differences in the territorial behaviour of the same individual according to the phases of territoriality (beginning, peak, end) which is shown by the example of Thomson's gazelle.
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This paper forms part of a general bio-ecological study of the tsessebe (Damaliscus lunatus lunatus Burchell) presently being undertaken in the Kruger National Park. Territorial demarcation and maintenance is achieved by various ritualized displays and marking techniques. Territory marking includes both visual and olfactory methods. Agonistic behaviour is evidenced by dominance and threat displays while territorial conflicts may result in actual fighting. The various behavioural patterns relating to territoriality are discussed with an evaluation of their possible significance in the social organisation of the species.
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The territorial behaviour of the black wildebeest was studied from 1968 to 1970 in the Willem Pretorius Game Reserve and other reserves in South Africa.Territoriality is a prerequisite for reproduction. Non-territorial males are barred from partaking in the rut. Territorial males may occupy a territory from their fourth year on, but the majority of territorial owners were five years old or more.Territorial males formed a network of territories, although single territorial males could also be observed. The spacing between individuals varied between 180-450 m in the Willem Pretorius Game Reserve; spacing of territorial black wildebeest as well as the number of territorial males depend on the density of the population.Territorial black wildebeest showed a strong and lasting attachment to their territories. The species was territorial throughout the year although absenteeism became prevalent after the rut.Territorial black wildebeest advertised their territories and defended them against intruding conspecific males. Advertising behaviour included demonstrative-threat advertising derived from marking as well as acoustic, static-optic and dynamic-optic advertising. The species did not demarcate a territory with either faeces or urine, or pre-orbital or interdigital glands.The encounters of territorial males were governed by highly ritualised motor patterns (Challenge Ritual). The ritual includes both aggressive and non-aggressive behaviour patterns.The Challenge Ritual ensures that only the fittest male can maintain his territory and thus partake in the rut. Furthermore, it serves to “satisfy“ the social needs of the solitary bulls in the highly gregarious species. Other functions of territorial behaviour include the separation of sexes and the spacing out of the population.The known agonistic and territorial behaviour of the various species of the tribe Alcelaphini is compared and discussed. The differences and similarities in behaviour do not follow the taxonomic separation of the various species but are indicative of different habitats.
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Three male and one female pronghorn (Antilocapra americana) were taken in the wild when a few hours to three days old and were hand-raised. The development of their behaviour was studied with particular emphasis on their olfactory communication. Fifty hours of field observations confirmed most of the findings in the captive animals.Four scents are important in the social life of the pronghorn: They originate with the paired subauricular and ischiadic glands, the single dorsal gland and the odour of urine. The subauricular and dorsal scents are produced by males only. Urine is an important component of the sniffing - pawing -urinating - defecation sequence of the males. The ontogeny of this sequence is described. An experiment with an artificial marking post demonstrated that the subauricular scent of a male one rank lower released the strongest response in other bucks. Preliminary gas chromatographic analysis of the subauricular scent has been carried out.
Chapter
Two wild members of the camel family occur on the aridlands of South America: the vicuna (Vicugna vicugna) and the guanaco (Lama guanicoe). The vicuna is found in the central Andean altiplano or puna, an equatorial short-grassland that is above treeline and below the snowline, despite its 3700m to 4800m elevation (Koford, 1957; Franklin, 1973 a, b). In contrast, the guanaco ranges from sea level to 4000m and is more widely distributed (Franklin, 1975).
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Publisher Summary This chapter discusses the information contained in the odorous secretions of mammals and provides a classification system based on the behavioral and chemical analyses. This classification divides social odors into two groups: identifier and emotive. Identifier odors are defined as those produced through the regular metabolic processes of the animal, without specific stimulation. The emotive odors are those produced as the result of some transient emotional state or external stimulus. The chapter categorizes nine different types of information contained in mammalian social odors: species, age, sex, colony membership, individuality, social status, reproductive state, maternal state, and stress odors. Social odors are modified by diet and hormone levels and by bacterial action. When the chemicals and bacteria responsible for producing the social odors have been identified, the responses of test animals show large individual differences. Responses to olfactory stimuli depend on hormonal and experiential factors. Theoretical models in the study of population regulation, sexual selection, kinship recognition, altruism, parental care, and territoriality infer that animals recognize particular individuals and specific relationships, and such recognition may depend to some extent on the information contained in olfactory signals.
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Territory, as here used, is meant to include an area or range exclusively occupied by an individual or group of individuals of a species, which is defined and recognized by them as their exclusive property or territory. Such a territory may be actively defended against invasion by others outside the territory of the individual group. Where not defended, the territory is nevertheless definitely marked by recognizable “sign posts” on the territory. What is perhaps more significant in regard to these sign posts is not so much the recognition of the territory by the animal or animals making the sign posts as the psychological effect that these sign posts have on non-members of the territory by an intimidation of the stranger entering such an occupied territory. Darling's (1937) explanation that home range is physiological and that territory is psychological is an attractively simple definition. However I doubt that it can be as simple and clear cut as this, and the author himself would probably be the first to admit that it would be extremely difficult to separate the physiological factors from the psychological factors that enter into the determination of either home range or territorial behaviorism. Territories are usually the direct or indirect result of breeding requirements or food needs or both. Territorialistic behavior is usually developed in direct relation to the social development of the animal, the greater the social development the greater the degree of territorialistic behavior. The sedentary habits and the tendency to occupy a well-defined home range is well recognized and generally understood in members of the Family Cervidae. Territorialism, on the other hand, apparently has been overlooked, or is not recognized in this group by most workers. Flerov (1952) and Kaplanov (1948) all give excellent descriptions of the rutting and general behavior of the various members …
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Otters were studied by reading tracks and signs in fresh water habitats in Southern Sweden, in 1958-1966. 1. The populations in winter consisted of 30-40% resident territory holders, about the same proportion of temporary residents or transients and 25-38% young of the year. The rate of reproduction of otters is low. Most probably some females do not breed every year. The density of otters in the areas studied was one otter per 0.7-1.0 km2 area of water or one individual per 2-3 km length of lake shore, and one otter per 5 km length of a stream. 2. Otters display territorial behaviour which is shown by signal activity, dispersion pattern and movements of the otters. 3. The adult dog otters maintain territories which have an individual character, their size and location depending on the qualities of the dogs. The boundaries are overlapping zones where territorial conflicts occur continuously. 4. The females with cubs (family groups) exploit territories separately. The areas are probably fixed and situated within the dog otters' territories. Territorial conflicts between the family groups are rare, after the areas have been settled. 5. Otters - apart from territory-holding dog otters and females with cubs - behave as temporary residents (mainly during summer and winter) or transients. 6. Territorial behaviour primarily appears between individuals of the same sex. The behaviour has a pronounced individual character. The individuals behave according to their status in a hierarchic system. 7. The territories are primarily maintained by threatening signals. Avoidance plays a larger part than pursuits. 8. The territories of the family groups are feeding areas, securing the young access of food all the first year of their lives. The dog otters' territories primarily have sexual significance. /// Исследовали популяции выдр в пресноводных водоемах Южной Швеции в 1958-1966 гг. 1. Популяции выдр зимой состоят на 30-40% из постоянных обитателей участков, примерно того же количества временных и кочующих обитателей и 25-38% молодых особей, родившихся в том же году. Скорость размножения у выдр очень низка. Некоторые самки очевидно не приносят потомства ежегодно. Плотность популяций выдр в исследованных участках составляла І экз. 0,7-1,0 KM2 водной поверхности или Іэкз. на 2-3 погонных км берега озера либо на 5 погонных км берега реки. 2. Выдры имеют индивидуальные участки, что доказывается характером их распределения и передвижения. 3. Взрослые самцы занимают определенные территории, являющиеся их индивидуальными участками. Территории соседних участков перекрываются у границ, и здесь постоянно возникают конфликты. 4. Самки с детенышами (семьн) занимают отдельные территории, расположенные внути индивидуальных участков. После заселения участков территориальные конфликты между семьями возникают редко. 5. Остальные представители популяции (исключая упомянутых выше самцов и самок с детенышами) ведут себя как временные или кочующие посетители участков, особенно летом и зимой. 6. Территориальная приуроченность проявляется сначала у представителей одного пола. Поведение выдр в отношении территории носит индивидуальный характер и зависит от положения каждого животного в иерархической системе. 7. Выдры охраняют свои участки от других особей, применяя сигналы угрозы. 8. Территории семейных групп - это места, хорошо обеспеченные пищей, где имеется возможность прокормить детенышей в течение первого года их жизни. Выбор участков взрослыми самцами связан с их половой активностью.
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This paper, covering the first 4 years of a continuing study, reports on preliminary findings concerning territoriality and its function in a mountain lion (Felis concolor) population. The research was carried out in the Idaho Primitive Area in central Idaho. Forty-three different lions were captured and marked during four winter and early spring seasons. Thirty-one individuals were recaptured 89 times, making a total of 132 captures during the study. Nine resident adults, captured a total of 59 times, provided the bulk of the data on home range and territoriality. Minimum size of the males' winter home range was constant from year to year, but it varied for females, depending upon their reproductive status. The smallest winter home range for a female, during a single season, was approximately 5 square miles, the largest approximately 20 square miles. Males utilized larger areas. Resident male lions occupied distinct winter territories without overlap, but resident females shared some common areas. Male territories overlapped those of females. Lions exhibited a high degree of tolerant but unsocial behavior. No evidence of territorial defense was noted. Transient lions of both sexes moved freely through occupied territories. A mutual avoidance behavioral mechanism acted to distribute lions in both time and space. Visual and olfactory marks serve to facilitate avoidance between lions.
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Conflicts between animals of the same species usually are of ``limited war'' type, not causing serious injury. This is often explained as due to group or species selection for behaviour benefiting the species rather than individuals. Game theory and computer simulation analyses show, however, that a ``limited war'' strategy benefits individual animals as well as the species.
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Marking behavior by thamins (or Eld's deer, Cervus eldi thamin) was observed in a zoo.
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The scent-marking behaviour of a group of captive but undisturbed muntjacs (Muntiacus reevesi) was studied for 450 hr in a 2200-m2 enclosure at the Calgary Zoo in 1971. Males marked more than females, and dominants of both sexes marked more than subordinates. Experimental alterations of the composition of the herd revealed that dominants marked most when in the presence of their subordinate rival. Some muntjacs covered the scent of conspecifics with their own. This behaviour was interpreted as a means of monitoring the circulation of conspecifics in one's home range.
Article
Marked Ss were investigated in Tanzania. Grouping patterns were either sedentary, with established territories, or nomadic, with temporary territories. Both patterns were interchangeable. Nomadism was a later adaptation. The individual distance was about 130 yds.; males became territorial at 34-40 mo., sometimes earlier. Since territory is a prerequisite to reproduction, aggressive competition dominated sexual behavior. Androgen level appeared to control both sex and aggression, but were linked to a fixed territory. Without a territory the gregarious behavior dominated and even mature males returned to the bachelor herd ("psychological castrates"). Territory owners constantly advertised their status, mainly through an elaborate "challenge ritual." (102 ref.) (PsycINFO Database Record (c) 2012 APA, all rights reserved)
Article
The eleven different functions for which mammals use urine marking are reviewed in this paper, and the urine marking behavior of the red fox (Vulpes vulpes) is described in detail. A new hypothesis is advanced that urine marking may serve as a "book keeping system" in the red fox's scavenging behavior. Foxes consistently investigate and urine mark inedible food remnants (e.g., bones, bird wings, and dried out pieces of hide). When a fox re-investigates a marked remnant, the urine mark signals "no food present," and the fox investigates this object for only a brief period of time. This use of urine marking may increase the efficiency of its scavenging behavior, i.e. more food-items found per hour of scavenging. This efficiency may be particularly important during periods of food shortage. The hypothesis is tested in three different experiments, using free-ranging red foxes as subjects. Experiment I establishes that fox do urine mark food remnants. Experiment II shows that foxes investigate for a significantly shorter period of time (P<0.001) food remnants exhibiting both the odor of food and the odor of urine as compared to remnants exhibiting just the odor of food. Experiment III suggests that there a hierarchy of stimuli which determines different responses in the fox's scavenging behavior. The experiments also suggest that there is a degree of social behavior in the scavenging activities of red foxes. Foxes appear to use each other's urine marks to increase the efficiency of their scavenging behavior. Thus this study definitely support LEYHAUSEN'S (1965) statement that the social life of solitary animals is frequently more complex than we realize. Solitary species probably show many ingeniously adapted mechanisms for occupying niches where highly social species could not be maintained. The social evolution and ecological advantages of solitary species deserve to be the focus of future research.
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Observations of wolves on Isle Royale are reported for 1961–66, with interpretations including the earlier 3-year period described by Mech (1966). On this 210-square-mile island the fully protected wolf population varied from approximately 22 to 28 in midwinter. The major and minor foods were moose and beaver, respectively. The main pack varied in number between 11 and 22 with about three breeding pairs believed present. The population remained relatively stable; mating occurred every winter; and adult mortality appeared to be low. High mortality among pups seemed to be the point of population control. Socio-economic factors may have controlled the size of the large pack. Availability of food during the period of parturition and rearing probably was critical to survival of young. Recruitment of young appeared to take place in years of high production of moose calves. Numbers in the large pack probably were curtailed through the progressive exclusion of aged and socially subordinate individuals. Under harassment these animals separated and became pack-following scavengers, then probably true loners ranging outside the area used by the pack. Smaller aggregations of two or three non-breeders were seen each winter, as were the loners, some of which appeared thin and weak. The only known breeding outside the big pack was in a group of five present in the winter of 1965. This group was probably a family unit which separated from the main pack. A year later the male had disappeared, and remains of a pup, probably theirs, were found. In the winter of 1966 the alpha male of the large pack became lame and apparently was killed. This background appears favorable for further changes in social organization.
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Scent marking by the beaver (Castor canadensis) has commonly been ascribed a function in territorial maintenance. This study examined the placement of scent mounds by beaver colonies, the frequency of scent mounds constructed by colonies with a low probability of encountering conspecifics versus colonies with a high probability of such encounters, and the responses of individual animals to castor gland homogenate from donor animals of known sex and age. Scent mounds are constructed most often in areas of high colony activity, and colonies with a high probability of encountering transient conspecifics construct significantly more mounds. Behavioral data suggest that information regarding sexual status may be transmitted by scent mounds. The data do not support a territorial maintenance function for scent marking in the beaver.
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In many large animals, changes in fighting ability within breeding seasons or across the lifetime of individuals are related to changes in body condition but not to obvious changes in size. In situations where a conflict of interests is likely to lead to a fight, we might consequently expect opponents to assess each other on traits which are related to variation in body condition. This appears to be the case among red deer stags. Competing stags engage in roaring ‘contests’ in situations where fights are likely. Observation combined with playback experiments showed that stags answered each others’ roars and that their roaring rate was related to that of their opponent. Both individual differences and temporal changes in roaring rates were correlated with changes in fighting ability and roaring contests usually occurred only where there was no obvious size discrepancy between opponents. The study suggested that assessment procedures probably continued during the parallel walks which commonly succeed roaring contests.