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Scent marking in shelter dogs: Effects of body size
... In a previous study of scent-marking behavior during first walks of dogs at the same shelter, we found that time spent at the shelter did not influence urinary behavior (frequency of urination or percent of urinations directed at targets in the environment) or likelihood of defecation or ground scratching [40]. In a subsequent study in which many dogs were walked multiple times [47], we found that time spent at the shelter positively influenced rate of urination, percent of directed urinations, and likelihood of defecation (ground scratching was not studied). In light of our findings for first walks [40], we interpreted the positive influence of time at shelter in our second study [47] as having resulted from our inclusion of multiple walks on individual dogs, and suggested that the positive influence of time spent at the shelter on marking behavior could reflect dogs becoming more familiar with their surroundings and routine, as well as with us [47]. ...
... In a subsequent study in which many dogs were walked multiple times [47], we found that time spent at the shelter positively influenced rate of urination, percent of directed urinations, and likelihood of defecation (ground scratching was not studied). In light of our findings for first walks [40], we interpreted the positive influence of time at shelter in our second study [47] as having resulted from our inclusion of multiple walks on individual dogs, and suggested that the positive influence of time spent at the shelter on marking behavior could reflect dogs becoming more familiar with their surroundings and routine, as well as with us [47]. For these two earlier studies [40,47], there were no single male walkers or single female walkers; we always had two people on each walk, one student (male or female) and B.M. (one person walked the dog and the other recorded behavioral observations). ...
... In a subsequent study in which many dogs were walked multiple times [47], we found that time spent at the shelter positively influenced rate of urination, percent of directed urinations, and likelihood of defecation (ground scratching was not studied). In light of our findings for first walks [40], we interpreted the positive influence of time at shelter in our second study [47] as having resulted from our inclusion of multiple walks on individual dogs, and suggested that the positive influence of time spent at the shelter on marking behavior could reflect dogs becoming more familiar with their surroundings and routine, as well as with us [47]. For these two earlier studies [40,47], there were no single male walkers or single female walkers; we always had two people on each walk, one student (male or female) and B.M. (one person walked the dog and the other recorded behavioral observations). ...
Interactions with humans influence the behavior and physiology of other animals, and the response can vary with sex and familiarity. Dogs in animal shelters face challenging conditions and although contact with humans typically reduces stress and behaviors associated with stress, evidence indicates that shelter dogs react differently to unfamiliar men and women. Given that some aspects of canine scent-marking behavior change under fearful conditions, we examined whether sex of an unfamiliar walker would influence scent-marking behavior of 100 shelter dogs during leash walks. Male dogs urinated at higher rates when walked by unfamiliar women than when walked by unfamiliar men; female dogs urinated at similar rates when walked by unfamiliar women and unfamiliar men. Sex of walker influenced urinary posture in male dogs, but not in female dogs. Both male and female dogs were more likely to defecate when walked by unfamiliar women than by unfamiliar men. Based on our findings that shelter dogs behave differently in the presence of unfamiliar men and women, we suggest that researchers conducting behavioral studies of dogs record, consider in analyses, and report the sex of observers and handlers as standard practice. We also recommend recording the sex of shelter staff present at behavioral evaluations because the results of these evaluations can impact dog welfare.
... For this observational study, I tested the hypothesis that gonadal hormones influence sexually dimorphic scent-marking behaviors in male dogs, but not female dogs (intact females were not in either proestrus or estrus). Urine-marking is highly dimorphic, with male dogs marking more frequently than female dogs (Beach, 1974;Cafazzo et al., 2012;McGuire, 2016;McGuire and Bemis, 2017). I predicted that gonadectomy would decrease urine-marking in males but have no effect on urine-marking in females. ...
... Although there is a tendency for more male than female dogs to display ground-scratching, significant sex differences in this behavior have not been reported (Bekoff, 1979;McGuire, 2016), so one would expect little or no effect of gonadectomy on ground-scratching by either males or females. Defecation is not a sexually dimorphic behavior in dogs (McGuire, 2016;McGuire and Bemis, 2017;Sprague and Anisko, 1973), so no effect of gonadectomy was predicted for defecation by either males or females. The hypothesis was tested using two different experimental designs. ...
... These findings of no difference between intact and spayed females with respect to likelihood of defecation at either shelter agree with an earlier report for privately owned female Jack Russell terriers (Wirant and McGuire, 2004). Other studies of scent-marking in dogs (McGuire, 2016;McGuire and Bemis, 2017;Sprague and Anisko, 1973) and coyotes (Gese and Ruff, 1997) reported no sex differences in frequency of defecation, which is consistent with what I found at the Tompkins shelter. In feral dog packs, however, placement of feces on conspicuous objects was limited to adult males of high social status (Cafazzo et al., 2012). ...
... In domestic dogs (Canis lupus familiaris), urination is the most common form of scent marking and urinary behavior varies greatly across sex, age, and size classes (Cafazzo, Natoli & Valsecchi, 2012;McGuire, 2016;McGuire & Bemis, 2017). Adult males typically use the raised-leg posture, juvenile males use the lean-forward posture during which no hindlimb is raised, and most females use the squat posture (Sprague & Anisko, 1973). ...
... Adult males urinate more frequently and direct more urinations onto vertically oriented targets than do adult females (McGuire, 2016). Small dogs urinate more frequently and direct more urinations than large dogs, which may indicate a preference by small dogs for communicating through scent marking rather than direct interactions that could be risky (McGuire & Bemis, 2017). ...
... Assuming that urine mark height communicates body size and corresponding competitive ability to conspecifics, Study 2 tested the hypothesis that urine marking is a dishonest signal in male dogs. Given that small dogs urinate more frequently and direct more urinations than large dogs (McGuire & Bemis, 2017) and would seem to benefit more from exaggerating their size, we predicted that small male dogs would exhibit larger raised-leg angles than large male dogs. ...
Scent marking is a common mode of communication in mammals. Such marking is thought to communicate information about the signaler's size and corresponding competitive ability and accurately reflect the signaler's attributes (i.e., an honest signal). However, new data suggest that scent marking can be dishonest in certain circumstances. Via two studies, we tested the hypothesis that urine marking is a dishonest signal in adult male domestic dogs, which raise a hindlimb when marking vertical objects. In Study 1, we tested whether raised-leg angle (i.e., during a urination, the angle between a dog's raised leg and the axis normal to the ground) is a proxy for urine mark height (n = 15 dogs) and, in Study 2, we tested whether small dogs exhibit larger raised-leg angles than large dogs (n = 45 dogs). We videotaped urinations of adult male dogs and, afterwards, measured height of urine marks (Study 1) and degree of raised-leg angles (Studies 1 and 2). In Study 1, we found significant positive relationships between both raised-leg angle and height of urine mark and body size (using either body mass or height at withers) and height of urine mark; raised-leg angle was a stronger predictor than either measure of body size. In Study 2, we found a significant negative relationship between body size (using either body mass or height at withers) and average raised-leg angle. Our findings support raised-leg angle as a proxy for urine mark height and provide additional evidence that scent marking can be dishonest. Assuming body size is a proxy for competitive ability, small adult male dogs may place urine marks higher, relative to their own body size, than larger adult male dogs to exaggerate their competitive ability. We did not control for over marking, which also may explain our findings.
... We assume that the role of scratch marks is correlated with mark height; if the mark is higher, the individual is larger and more competitive, similar to AGS and urine marks (White et al., 2002); however, the specific role of scratch marks needs to be further investigated. McGuire and Bemis (2017) found that body size affects the frequency of marking in a study of domestic dogs (Canis lupus familiaris; Mcguire & Bemis, 2017). However, from studies of pandas, we know that only body size reflects their competitive ability (Nie, Swaisgood, Zhang, Liu, & Wei, 2012), but whether it affects the frequency of marking deserves further investigation. ...
... We assume that the role of scratch marks is correlated with mark height; if the mark is higher, the individual is larger and more competitive, similar to AGS and urine marks (White et al., 2002); however, the specific role of scratch marks needs to be further investigated. McGuire and Bemis (2017) found that body size affects the frequency of marking in a study of domestic dogs (Canis lupus familiaris; Mcguire & Bemis, 2017). However, from studies of pandas, we know that only body size reflects their competitive ability (Nie, Swaisgood, Zhang, Liu, & Wei, 2012), but whether it affects the frequency of marking deserves further investigation. ...
Inter‐animal communication allows signals released by an animal to be perceived by others. Scent‐marking is the primary mode of such communication in giant pandas ( Ailuropoda melanoleuca ). Signal detection theory propounds that animals choose the substrate and location of their scent marks so that the signals released are transmitted more widely and last longer. We believe that pandas trade‐off scent‐marking because they are an energetically marginal species and it is costly to generate and mark chemical signals. Existing studies only indicate where pandas mark more frequently, but their selection preferences remain unknown. This study investigates whether the marking behavior of pandas is consistent with signal detection theory. Feces count, reflecting habitat use intensity, was combined with mark count to determine the selection preference for marking. The results showed that pandas preferred to mark ridges with animal trails and that most marked tree species were locally dominant. In addition, marked plots and species were selected for lower energy consumption and a higher chance of being detected. Over 90% of the marks used were the longest‐surviving anogenital gland secretion marks, and over 80% of the marks were oriented toward animal trails. Our research demonstrates that pandas go out of their way to make sure their marks are found. This study not only sheds light on the mechanisms of scent‐marking by pandas but also guides us toward more precise conservation of the panda habitat.
... We assume that the role of scratch marks is correlated with mark height; if the mark is higher, the individual is larger and more competitive, similar to AGS and urine marks (White et al., 2002); however, the specific role of scratch marks needs to be further investigated. McGuire and Bemis (2017) found that body size affects the frequency of marking in a study of domestic dogs (Canis lupus familiaris; Mcguire & Bemis, 2017). However, from studies of pandas, we know that only body size reflects their competitive ability (Nie, Swaisgood, Zhang, Liu, & Wei, 2012), but whether it affects the frequency of marking deserves further investigation. ...
... We assume that the role of scratch marks is correlated with mark height; if the mark is higher, the individual is larger and more competitive, similar to AGS and urine marks (White et al., 2002); however, the specific role of scratch marks needs to be further investigated. McGuire and Bemis (2017) found that body size affects the frequency of marking in a study of domestic dogs (Canis lupus familiaris; Mcguire & Bemis, 2017). However, from studies of pandas, we know that only body size reflects their competitive ability (Nie, Swaisgood, Zhang, Liu, & Wei, 2012), but whether it affects the frequency of marking deserves further investigation. ...
The purpose of inter-animal communication is to allow signals released by the animal to be perceived by others. Scent marking, with its characteristics of specificity and delay, is thought to be the primary mode of communication in giant pandas (Ailuropoda melanoleuca). The “signal detection theory” predicts that animals choose the substrate and location of their scent marks so that the signals released are transmitted more widely and remain longer. As an energetically marginal species, the cost of generating and marking chemical signals is costly for pandas, so they are predicted to make trade-offs in scent marking. However, existing studies do not account for the selective preferences of pandas for marking, as they are only explained by the density of marks at a certain location. Our study wanted to investigate whether the marking behavior of pandas is indeed consistent with signal detection theory. For the first time, we propose to use fecal counts to reflect the intensity of habitat use by pandas, combined with mark counts to determine the selective preference for marking. Our findings show that the scent marking behavior of pandas is consistent with signal detection theory, and that they go out of their way to ensure that their marks are detected. The results of the study will help us to further develop the conservation of pandas and their habitats.
... However, even without direct contact, dogs can communicate with each other through chemical signals [66]. Urine-marking, which involves leaving urine or other body secretions on distinctive elements of the landscape [67], is the most common form of scent marking [68]. The marked object, such as a tree or a streetlamp, becomes a scent message containing much information about the individual who left it [65]. ...
... Male dogs and wolves mark more frequently in unfamiliar areas and will continue to do so even if their bladder is empty, implying that marking is not associated with urine passage [70]. Males urinate more frequently and direct more urinations onto vertically oriented targets than females [71], and small dogs urinate more frequently than large dogs, probably because scent marking involves less risk than direct interaction [68]. ...
Olfaction in dogs is crucial for gathering important information about the environment, recognizing individuals, making decisions, and learning. It is far more specialized and sensitive than humans’ sense of smell. Using the strength of dogs’ sense of smell, humans work with dogs for the recognition of different odors, with a precision far exceeding the analytical capabilities of most modern instruments. Due to their extremely sensitive sense of smell, dogs could be used as modern, super-sensitive mobile area scanners, detecting specific chemical signals in real time in various environments outside the laboratory, and then tracking the odor of dynamic targets to their source, also in crowded places. Recent studies show that dogs can detect not only specific scents of drugs or explosives, but also changes in emotions as well as in human cell metabolism during various illnesses, including COVID-19 infection. Here, we provide an overview of canine olfaction, discussing aspects connected with anatomy, physiology, behavioral aspects of sniffing, and factors influencing the olfactory abilities of the domestic dog (Canis familiaris).
... It can, however, also act as a display of dominance or social challenge among both wolves and dogs (Asa et al., 1990;Cafazzo et al., 2012). There is some evidence that scent marking can even indirectly communicate information like body size without direct interaction (McGuire & Bemis, 2017;McGuire et al., 2018). ...
... All-female dyads displayed no urination and only one of the 12 dogs, which displayed this behavior, was female. This squares with other findings showing that males engage more frequently in marking behavior (Beach, 1974;Bekoff, 1979;Howse et al., 2018;McGuire & Bemis, 2017;Pal, 2003). ...
Introduction:
The aim of this study was to explore spontaneous social interactions between dyads of unfamiliar adult dogs. Although intraspecific encounters are frequent events in the life of pet dogs, the factors that might influence encounters, such as sex, dyad composition, reproductive status, age, and state of cohabitation (keeping the dogs singly or in groups), remained unexplored.
Methods:
In this study, we assigned unfamiliar, non-aggressive dogs to three types of dyads defined by sex and size. We observed their unrestrained, spontaneous behaviors in an unfamiliar dog park, where only the two dogs, the owners, and experimenter were present.
Results:
We found that the dogs, on average, spent only 17% of the time (less than 1 min) in proximity. Sex, dyad composition, reproductive status, and age influenced different aspects of the interactions in dyads. Female dogs were more likely to initiate the first contact in their dyad but later approached the partner less frequently, were less likely to move apart, and displayed less scent marking. Following and moving apart were more frequent in male-male interactions. Neutered dogs spent more time following the other dog and sniffed other dogs more frequently. The time companion dogs spent in proximity and number of approaches decreased with age.
Conclusion:
The study provides guidance for dog owners about the outcomes of intraspecific encounters based on the dog's age, sex, and reproductive status, as well as the sex of the interacting partner.
... The data presented in this article were collected between February 16, 2013 and December 9, 2016, as part of a long-term research effort focused on scent-marking behavior of shelter dogs. Details of care and housing of dogs and our methods for classifying dogs by size class and age class have been presented elsewhere (Gough and McGuire, 2015;McGuire 2016;McGuire and Bemis, 2017); therefore, we provide brief descriptions here. ...
... Body size influenced urinary posture in male and female shelter dogs in a manner consistent with the emerging picture of small dogs as preferentially using urine-marking as a means of communication (McGuire and Bemis, 2017). Time spent at shelters influenced urinary posture in adult and senior males, but not females, with likelihood of using the raised-leg posture increasing with time. ...
Motor laterality is the preferential use of structures on one side of the body. Although domestic dogs are commonly used in laterality research, few studies have examined hindlimb motor tasks, and no study, to our knowledge, has examined the effects of body size on measures of laterality. First, we observed 659 dogs during walks at 2 animal shelters to assess likelihood of raising a hindlimb during urination in relation to body size, age class (juvenile, adult, and senior), reproductive status (intact vs. spayed or neutered), shelter, and time spent at shelters; males and females were analyzed separately, and we excluded juveniles from statistical analyses because no female in this age class raised a hindlimb. Second, we observed a subset of the adult and senior dogs (n = 46) over multiple walks to determine if they exhibited side preferences with respect to hindlimb raised during urination. We looked for individual biases and a population bias and analyzed hindlimb preferences with respect to body size. For both males and females, small dogs were more likely than medium and large dogs to raise a hindlimb during urination. Intact males (96%) were more likely than neutered males (83%) to have a predominant urinary posture with a raised hindlimb. Time spent at the shelters influenced urinary posture in mature males: with each additional day spent at the shelter, the odds of a male having a predominant urinary posture with a raised hindlimb increased by 2%. We found the following distribution of hindlimb preferences in the subset of dogs studied for laterality: right preferent, 19.6%; left preferent, 30.4%; and ambilateral, 50.0%. There was no evidence of a population-wide bias for hindlimb raised. Body size did not influence laterality strength or whether a dog was hindlimb preferent (right or left) versus ambilateral. Thus, the likelihood of raising a hindlimb when urinating, but not laterality with respect to hindlimb raised, can be added to the growing list of behavioral differences between small dogs and larger dogs. Previous studies have reported that under stressful conditions, mature male dogs may temporarily revert to the juvenile urinary posture in which no hindlimb is raised. Our finding for adult and senior males that likelihood of raising a hindlimb during urination increased with time spent at the shelters suggests that male urinary posture could potentially serve as a behavioral indicator of adjustment to shelter conditions.
... Indeed, the domestic dog can transmit information to conspecifics relating to individual identity, including sex and social status. One method which dogs use to diffuse chemical signals is urine marking (Catala et al. 2019;McGuire Bemis 2017;Horowitz 2017;Cafazzo et al. 2012) as illustrated in figure 4. In female dogs, urine marking usually occurs around oestrus, indicating it is a signal related to mate attraction and sexual reproduction. However, urine marking in males, and also, to a lesser extent, females, is thought to be related to territory defence and dominance hierarchies (Cafazzo et al. 2012). ...
Olfaction, as a semiotic modality, receives relatively less attention than other sensory modalities. However, chemiosemiosis and semiochemicals are fundamental components of zoosemiosis, occurring across animal taxonomic groups. Indeed, olfaction is thought to be one of the most ancient sensory modalities from an evolutionary perspective and significantly, even unicellular organisms, such as the bacterium Escherichia coli , utilize a form of chemiosemiosis when foraging for nutrients, as part of a process known as ‘chemotaxis’. Further, many taxonomic groups have evolved to produce dedicated ‘semiochemicals’ (often known as pheromones or allomones) which have the sole purpose of being diffused into the environment as a social signal. In this paper, I highlight the importance of Umwelt theory when studying animal communication, by reviewing the less conspicuous and intuitive chemiosemiotic modality, across animal taxa. I then go on to discuss chemiosemiosis within a linguistic framework and argue that complex pattern recognition underpins linguistic theory. Thus, I explore the concept that chemiosemiosis has features in common with language, when the factor of time, in the transmission and decoding of a signal, is taken into account. Moreover, I provide discursive evidence in support of a unified theory of sensory perception, based on structural and functional aspects of signal transmission and cognitive complex pattern recognition. I conclude by proposing a chemosemiotic hypothesis of language evolution.
... 2012), there exist well-known examples pointing to the importance of chemical communication in a wide range of domestic species and biological contexts that could have implications for horse management and welfare. These include studies on the identification of reproductive status and sexual receptivity in females (Goodwin et al., 1979;McGlone and Morrow 1987;Burger et al., 2018), mother-young recognition (Lévy et al., 2004), territorial marking (Bradshaw and Cameron-Beaumont, 2000;McGuire and Bemis, 2017), and discrimination of group members from unfamiliar conspecifics (Mykytowycz, 1962;Perón et al., 2014). Furthermore, the sense of smell plays an important role in human-horse interactions; for example, human body odorants may induce sympathetic and parasympathetic changes and stimulate horses emotionally (Lanata et al., 2018). ...
This work provides an overview of the role of chemical communication in horse behavior, and addresses the potential usefulness of this knowledge for management practices and animal welfare. First, we present an outline of the social organization of horses under natural conditions, and the problems that may emerge due to domestication. Second, we briefly describe the mammalian olfactory system, noting the peculiarities of horses. Third, we describe some behaviors in which horses use chemical communication, including sexual behavior, and mother-young relationship. Finally, the article focuses on the gaps in information on the chemical communication of the horse, and the underexplored possible advantages of using chemical signals to reduce the stress response of foals and horses in the context of domestication.
... Previous research has focused on the development of sex differences in canine urine marking behavior but only our previous preliminary research has evaluated the relationship between marking and urinary cortisol levels [23]. More recently, body size was reported to influence urine marking frequency in shelter dogs with the hypothesis that small dogs preferentially communicate via scent marking because it is an indirect mode of communication that allows them to avoid costly direct interactions [40]. The weak positive correlation between C/Cr and frequency of urine marking could be related to the increased release of aldosterone under ACTH stimulus or to chemical information that dogs are in a stressful situation. ...
The aim of this study was to evaluate the effects of intraspecific social exposure (socialization vs rest) and habituation factors on the levels of urinary serotonin and cortisol [corrected respectively for creatinine to give the serotonin-creatinine ratio (5-HT/Cr) and cortisol-creatinine ratio (C/Cr)] and how they fluctuate in dogs. The frequency of marking during social sessions was recorded to evaluate its relationship with physiological parameters. The effects of covariates on 5-HT/Cr and C/Cr were assessed using a linear mixed models. 5-HT/Cr values were higher at dawn than at dusk during resting days, however, this difference is less evident after socialization sessions. During rest days, there was a trend for a decrease in C/Cr between dawn and dusk, while during social exposure days there was an opposite trend. Significant interactions were found between social exposure vs rest (P = 0.0005) and social exposure vs sessions (P = 0.002). Urine marking was more frequent in male than female dogs. The frequency of urine marking also appeared to be positively associated with C/Cr. Non-invasive monitoring of physiological markers could be a useful tool in assessing behaviour modulation following intraspecific socialization exposure. C/Cr was characterized by high individual variability and interactions with examined factors. The identification of new markers such as serotonin for assessing welfare in dogs is highly desirable.
Dogs and cats are popular companion animals that live together with humans. This special issue provides topics about the scents and olfaction of dogs and cats from the viewpoint of ethology. The first half introduces the significance of odors of feces, urine, and body, which is easily perceived as unpleasant odors, and how to deal with them. The second half mainly introduces the olfactory ability, the application of “Nosework” to animal welfare, and the effects of the owner’s body odor on the attachment behavior, in dogs and cats.
High demand for dogs in countries like the UK can lead to illegal intensive breeding and illegal importation of puppies for the pet trade. The current study investigates the effects of intensive breeding or ‘puppy farming’ on canine behaviour, explores new ways of predicting negative outcomes and categorising dog behaviour, and probes whether various types of training or routines can mitigate these behavioural outcomes. Participants completed an online self-report questionnaire, combining a shortened version of the Canine Behavioural Assessment and Research Questionnaire (mini C-BARQ) (Duffy et al., 2014), with new scales created in collaboration with the Scottish Society for the Prevention of Cruelty to Animals (Scottish SPCA). 2026 participants completed the questionnaire; most owners had dogs from non-puppy farm backgrounds (n = 1702), the rest had dogs from puppy farms (n = 123), or were unsure of the source of the dog (n = 201). We validated the mini C-BARQ as a tool for measuring dog behaviour, and explored its latent dimensions using factor analysis, extracting five first-order factors and one overarching second-order factor. We also confirm the validity of three of the four new scales developed with Scottish SPCA used to measure the impact of puppy farming practices. Linear and logistic regressions demonstrated that dogs from puppy farms have less desirable behaviours than dogs from other sources on 11 of the 14 behavioural subscales of the mini C-BARQ (for significant subscales, coefficients were between 0.1 and 0.2, and odds Ratios between 1.6 and 2.5). Generalized Linear Models (GLM) revealed the predictive power of two newly developed scales measuring early life experience in explaining variations in dog behaviour. In a GLM accounting for the dog's early life experience (and controlling for variables like breed and age), dog-walking significantly reduced the incidence of undesirable behaviours (p < 0.001), while different types of training had a significant interaction with poor early life experience in moderating canine behaviour (p < 0.002). Finally, dogs from puppy farms had significantly worse medical scores than dogs from other breeding sources (U = 144,719, z = 7.228, p < 0.001). These results suggest that puppy farming has negative impacts on dog behaviours and health, while more research is necessary to fully explore how to mitigate the effects of poor early life experience.
Scent marking is an important means of communication in mammals and many species elevate their
scent marks by depositing them on vertical objects. Traditionally, it has been assumed that marks are
elevated to increase detectability, but elevated marking may have evolved to convey information about
depositor size. Because the height of an elevated mark is typically dictated by the height of the marker,
receivers could use the physical location of an elevated scent deposit to indirectly assess the size, and
hence the competitive ability, of rivals. Considering that intrasexual competition is one of the primary
motivators of mammalian scent marking, and that the ability to assess a rival indirectly would provide
very real benefits (eliminating the need for potentially dangerous encounters), it would be surprising if
species were not utilizing mark height in this way. Nevertheless, it remains unknown whether any
mammal extracts such information from elevated scent marks. I tested whether wild dwarf mongooses,
Helogale parvula, discriminate between handstand scent deposits of differing height. Handstand marking
is an extreme form of elevated marking, with the marker balancing on its forepaws while flinging its hind
legs into the air and smearing anogenital secretions one full body length above the ground. I found that
females spent twice as long investigating anal-gland deposits positioned 16 cm above the ground, as
compared with those at 10 cm, even though the two were swipes of the same scent deposit and presumably
did not differ chemically. A faeces presentation experiment showed that females were more
interested in obtaining information about same-sex than opposite-sex conspecifics. I suggest that female
dwarf mongooses, which experience extreme intrasexual competition, use the height of handstand scent
marks as an indicator of depositor size, allowing them to concentrate their information gathering on
same-sex rivals that pose the greatest threat.
Motor laterality is the preference shown for using one limb or lateral half of the body over the other. In domestic dogs, most laterality studies have examined forelimb preferences during staged tasks. We focused instead on hindlimb preferences during urination when males use the raised-leg posture and females the squat-raise. We observed individual dogs during walks at two shelters (Tompkins County SPCA and Cortland Community SPCA) and recorded posture used for each urination and hindlimb raised, if any. First, we examined whether raising a hindlimb during urination varied with sex, age class, or reproductive status (females, anestrous intact or spayed; males, intact or neutered). Second, for dogs that raised a hindlimb during urination, we determined whether a population bias existed. Finally, for dogs with at least 10 urinations in which a hindlimb was raised, we examined whether a significant hindlimb preference existed. For some analyses, we had sufficient dogs at only one shelter. We found that males were more likely than females to raise a hindlimb during urination (P < 0.0001 at each shelter), and that propensity to raise a hindlimb was unaffected by reproductive status (P = 0.82, Cortland). Seniors were more likely than adults, which, in turn, were more likely than juveniles, to raise a hindlimb during urination (P < 0.0001, Tompkins). We found no evidence of a population bias with respect to hindlimb raised at either shelter (% of hindlimb raises involving the right hindlimb: 53%, Tompkins; 43%, Cortland). Of the dogs that met the criterion for at least 10 urinations with a raised hindlimb, most were ambilateral (83%, Tompkins; 90%, Cortland). Our study confirms and extends for shelter dogs the effects of sex and age on urinary postures previously reported for dogs living under other conditions; to our knowledge, the increased likelihood of raising a hindlimb during urination that characterized seniors (males, 91%; females, 25%) when compared to adults (males, 73%; females, 6%) has not been reported previously. Lack of a population bias with respect to hindlimb raised is consistent with findings of most motor laterality studies in dogs. However, our finding that most dogs were ambilateral differs from results obtained from studies using staged forelimb tasks. Assessing motor laterality for a natural hindlimb behavior in dogs during walks has both advantages and disadvantages, which include ease of observation during a positive experience for the dog and the challenge of obtaining sufficient scores for each dog.
Most mammals scent‐mark, and a variety of hypotheses have been put forward to explain this behaviour. Most of our knowledge about scent marking in domestic dogs comes from studies carried out on laboratory or companion dogs, while few studies have been carried out on free‐ranging dogs. Here, we explored the functional significance of different scent‐marking behavioural patterns in a pack of free‐ranging domestic dogs by testing two non‐exclusive hypotheses: the indirect territorial defence and the dominance/threat hypotheses. Through direct observation, we recorded the locations of dog scent marks (urination, defecation and ground scratching) and information regarding the identity and posture of the marking animal. We found evidence that markings are used by dogs to form a ‘property line’ and to threaten rivals during agonistic conflicts. Both males and females utilized scent marking to assert dominance and probably to relocate food or maintain possession over it. Raised‐leg urination and ground scratching probably play a role in olfactory and visual communication in both males and females. Urinations released by females, especially through flexed‐leg posture, may also convey information about their reproductive state. Finally, our observations suggest that defecation does not play an essential role in olfactory communication among free‐ranging dogs and that standing and squat postures are associated with normal excretion. Our results suggest that many of the proposed functions of marking behaviours are not mutually exclusive, and all should be explored through detailed field and laboratory studies.
Many mammal species adopt marking postures that elevate their scent deposits. The most extreme of these is handstand marking, in which an individual reverses against an upright object, flings its hind legs into the air above its back and balances bipedally on its fore feet. The resulting anogenital deposit is thus raised one full body length above ground level. It has been suggested that this energetically costly form of marking serves to provide conspecifics with information about the marker's body size and hence competitive ability. However, this explanation assumes that the height of an individuals’ deposit does reflect accurately its body size, an assumption that has never been tested in any hand‐standing species. This study investigated the relationship between body size and handstand mark height in a wild population of dwarf mongooses (Helogale parvula) in South Africa. We found that although body size and marking height were correlated positively for female dwarf mongooses, they were not related for males. Male dwarf mongooses (who are subject to intrasexual competition from outside their group) invested more heavily in anogenital range marking, marking at three times the female frequency and placing their deposits significantly higher than females (although they were not dimorphic). Males that were particularly vulnerable to rivals (i.e. those that were small for their age) tended to mark higher than more robust age‐mates, in keeping with the predictions of Adams & Mesterton‐Gibbons’ (1995, J. Theor. Biol.175, 405–421). model of deceptive threat communication. These findings suggest strongly that the height of anogenital scent deposits is of social significance to dwarf mongooses.
Dogs offer unique opportunities to study correlations between morphology and behavior because skull shapes and body shape are so diverse among breeds. Several studies have shown relationships between canine cephalic index (CI: the ratio of skull width to skull length) and neural architecture. Data on the CI of adult, show-quality dogs (six males and six females) were sourced in Australia along with existing data on the breeds' height, bodyweight and related to data on 36 behavioral traits of companion dogs (n = 8,301) of various common breeds (n = 49) collected internationally using the Canine Behavioral Assessment and Research Questionnaire (C-BARQ). Stepwise backward elimination regressions revealed that, across the breeds, 33 behavioral traits all but one of which are undesirable in companion animals correlated with either height alone (n = 14), bodyweight alone (n = 5), CI alone (n = 3), bodyweight-and-skull shape combined (n = 2), height-and-skull shape combined (n = 3) or height-and-bodyweight combined (n = 6). For example, breed average height showed strongly significant inverse relationships (p
To investigate the social roles of countermarking in dogs, we measured tail base position (TBP, a measure of social status), adjacent-marking and overmarking responses of male, nonoestrous female, gonadectomized male and gonadectomized female dogs to controlled presentations of urine from unfamiliar dogs and social groupmates. We also recorded dog sex, TBP, countermarking and urine investigation at a park. In urine presentations, only males overmarked, intact males (but not gonadectomized males) preferentially overmarked intact female urine, overmarking males had higher TBP than males that did not countermark and urine familiarity did not affect overmarking. In contrast, dogs adjacent-marked only unfamiliar samples, and neither sex nor TBP significantly affected adjacent marking. Gonadectomy did not significantly change the likelihood of countermarking. In dog park observations, males and females marked at and investigated ‘scent posts’ comprised of serial countermarks, often associated with visual landmarks. Males and females were equally likely to countermark and investigate urine and countermarks made up a similarly large portion of countermarking for males and females. Males and females with higher TBP urinated, investigated urine, and countermarked more than same-sex dogs with lower TBP. These studies suggest that although intact males may overmark female urine in part to guard mates as previously hypothesized, both sexes, intact and gonadectomized, likely countermark competitively. Social and sexual patterns also suggest that overmarks and adjacent marks may be distinct signals.
Behavioral evaluations of 2017 shelter dogs were used for identifying dogs with aggressive tendencies and for predicting post-adoption behavior problems. Associations between failure of the behavioral evaluation and demographic factors (age, breed, and sex) and the dog's behavioral history, evaluated by logistic regression, were highly significant (P72 months). Dogs that failed the behavior evaluation were not placed for adoption; therefore it was not possible to study prospectively the capability of behavioral evaluation to predict future aggressiveness in these dogs. Instead we developed tests for classifying dogs as aggressive or not aggressive based on their demographic factors and behavior evaluation outcomes. The results were compared retrospectively to the dogs’ known behavioral histories, which were obtained at intake to the shelter. This allowed estimation of the sensitivity, specificity, and accuracy of the classification tests. The most significant postdictor (i.e., “retrospective predictor”) of aggressiveness was failure of the behavioral evaluation (odds ratio 11.83, P
The urinary behavior of adult domestic dogs (Canis familiaris) is sexually dimorphic with respect to the posture (males lift a leg and females squat), frequency of urination (males urinate more frequently than females), and tendency to direct urine at specific objects in the environment (males are more likely than females to direct their urine). Such behavioral differences have led to the belief that urination functions largely, or exclusively, in elimination in female dogs, while having the additional function of scent marking in male dogs. In this study, we observed urinary behavior of six spayed and six non-estrous intact female Jack Russell Terriers during walks on and off their home area. The females ranged in age from 0.4 to 11.2 years. Frequency of urination was positively correlated with age, and females four or more years old directed the majority of their urinations at objects in the environment. Overall, females urinated more frequently and directed more of their urinations when walked off their home area than when walked within their home area. Spayed females were more likely than non-estrous intact females to ground-scratch following defecation; we detected a similar trend for ground-scratching after urination. There was, however, considerable variation among spayed females in the tendency to display ground-scratching behavior. Overall, the most common posture displayed by females while urinating was the squat-raise. Other postures, in order of their frequency of occurrence included squat, arch-raise, combination, and handstand. Females used the squat-raise and arch-raise postures more when off their home area than when on their home area. Overall, there was substantial individual variation among females in the postures used while urinating. Our data indicate that female urinary behavior varies with location and reproductive status, and that substantial individual differences exist among females for some patterns of behavior. Additionally, the large percentages of directed urinations by spayed (60.8%) and non-estrous intact females (56.7%) in our study suggest that urination in female dogs does not function solely in elimination, but that it also has a significant role in scent marking, even when females are not in estrus.
Scent-marking is a ubiquitous form of olfactory signaling in male mammals and both territorial males in resource-defense mating systems and dominant males in dominance mating systems scent-mark. A large body of evidence suggests a link between scent-marking by male mammals and intrasexual competition. Resource holders appear to mark to help establish and maintain their status. They may do this because scent marks allow potential opponents to assess the status or RHP of the signaler. Nonresource holding competitors benefit because they can adjust the level of escalation in relation to potential costs and benefits and avoid risky contests. Resource holders benefit through reduced costs because many nonresource holders withdraw to avoid escalated contests.
Large body size confers a competitive advantage in animal contests but does not always determine the outcome. Here we explore the trade-off between shortterm achievement of high social status and longer-term life history costs in animals which vary in competitive ability. Using laboratory mice, Mus musculus, as a model system, we show that small competitors can initially maintain dominance over larger males by increasing investment in olfactory status signalling (scent-marking), but only at the cost of reduced growth rate and body size. As a result they become more vulnerable to dominance reversals later in life. Our results also provide the first empirical information about life history costs of olfactory status signals.
Plasmacortisol levels were examined to assess the stress of dogs in a county animal shelter. Groups of dogs confined in the shelter for their 1st, 2nd, or 3rd day had higher cortisol levels than did a group maintained in the shelter for more than 9 days. Dogs in the shelter for an intermediate period (Day 4-9) had intermediate levels of cortisol. The cortisol concentrations of dogs during their first day in the shelter were greater than either those of the same dogs on Day 4/5 in the shelter or those of a group of pet dogs sampled in their own homes. There was no overall effect of 20 min of social interaction with a human (e.g., petting) on the plasma cortisol levels of dogs in the shelter on Day 1-3. However, the gender of the petter did affect cortisol levels. Those dogs interacting with a female had lower cortisol concentrations at the end of the session than did dogs interacting with a male. The results suggest that confinement in a public animal shelter produces a prolonged activation of the hypothalamic-pituitary-adrenal axis. Further, it appears that some subtle aspect of interaction with a human may be capable of moderating this response. Possible implications for the welfare of confined dogs, and for the development of behavior problems in dogs obtained from shelters, are discussed.
Behavior and plasma cortisol levels were examined in puppies and juvenile/adult dogs admitted to a public animal shelter. A behavioral test was developed to assess the responses of the dogs to novel or threatening conditions. Factor analysis of the behavioral responses of 166 dogs on day 3 in the shelter yielded six factors (locomotor activity, flight, sociability, timidity, solicitation, and wariness) that accounted for 68% of the total variance. Among those dogs remaining in the shelter for 9 days, plasma cortisol levels declined from day 2 to 9. Cortisol levels were weakly related to factor scores. In order to explore the relation of measures in the shelter to later behavior, questionnaires assessing problem behaviors were mailed to new owners of dogs 2 weeks and 6 months following adoption. Among puppies, wariness scores were negatively correlated with behavior problems at 2 weeks and cortisol levels were negatively correlated with behavior problems at 6 months. These results suggest how measures of behavior and endocrine activity obtained in shelters might prove useful for screening dogs for adoption or targeting dogs for behavioral intervention.
In this study, 60 shelter dogs (Canis familiaris) were observed in the modified version of the Strange Situation Test, which has proved to be a useful method for studying dogs' attachment behavior toward humans (J. Topál, A. Miklósi, V. Csányi, & A. Dóka, 1998). Before testing, 40 dogs were handled 3 times for 10 min. In the test, handled dogs encountered 2 persons: the handler in the role of the "owner" (OW) and an unfamiliar person (UP), whereas the 20 nonhandled dogs encountered unfamiliar persons in both roles. Dogs in the handled group exhibited more contact seeking with the entering OW, less physical contact with the UP, less frequent following of the leaving UP, and less standing by the door in the presence of the OW. The specific response of the handled dogs toward the handler fulfilled the operational criteria of attachment. In shelter conditions, the remarkable demand for social contact with humans may result in rather fast forming of attachment even in adult dogs.
Development of scent-marking behavior from the neonatal period to early adulthood has been well studied in dogs, but there is a distinct lack of information for older dogs. I studied scent-marking behavior during single walks of 500 male and female dogs at two shelters (Tompkins County SPCA and Cortland Community SPCA). My sample included juveniles, adults, and seniors. I found a significant effect of age on frequency of urination (_P_ < 0.0001 at both shelters): seniors urinated more frequently than adults (contrast significant at the Cortland shelter; _P_ < 0.07 at the Tompkins shelter), which urinated more frequently than juveniles. Age also influenced likelihood of directing urinations at targets in the environment (_P_ < 0.0001 at both shelters): seniors directed more of their urinations than did adults (significant at Cortland shelter only), which directed more of their urinations than did juveniles. I found that males urinated more frequently than females (_P_ < 0.0001 at both shelters) and directed more of their urinations (_P_ < 0.0001 at both shelters). Significant age and sex differences did not characterize defecation at either shelter. Ground scratching, whether after urination or defecation, was rarely performed by juveniles (% that ground scratched at least once: Tompkins shelter, < 14%; Cortland shelter, 0%), so I excluded juveniles from analyses of this behavior. Ground scratching after urination was not associated with sex or age (adults versus seniors) at either shelter, but was positively associated with number of directed urinations (Tompkins shelter, _P_ < 0.0001; Cortland shelter, _P_ < 0.002). Ground scratching after defecation was not associated with sex at either shelter, but was associated with age at the Tompkins shelter (_P_ < 0.03; % that ground scratched after at least one defecation: 28% of adults; 42% of seniors); a similar pattern occurred at the Cortland shelter (29% of adults; 50% of seniors), but the association failed to reach statistical significance perhaps due to smaller sample sizes. Finally, at the Tompkins shelter, ground scratching after defecation was positively associated with number of urinations followed by ground scratching (_P_ < 0.0001); here, again, a similar pattern occurred at the Cortland shelter but the association failed to reach statistical significance. These data reveal new relationships between scent-marking behaviors; indicate that some marking behaviors continue to change even after a dog has reached adulthood; and highlight the differential effects of sex and age on urination, defecation, and ground scratching.
Many mammals use scent marks to advertise territory ownership, but only recently have we started to understand the complexity of these scent signals and the types of information that they convey. Whilst attention has generally focused on volatile odorants as the main information molecules in scents, studies of the house mouse have now defined a role for a family of proteins termed major urinary proteins (MUPs) which are, of course, involatile. MUPs bind male signalling volatiles and control their release from scent marks. These proteins are also highly polymorphic and the pattern of polymorphic variants provides a stable ownership signal that communicates genome-derived information on the individual identity of the scent owner. Here we review the interaction between the chemical basis of mouse scents and the dynamics of their competitive scent marking behaviour, demonstrating how it is possible to provide reliable signals of the competitive ability and identity of individual males. (C) 2004 Wiley Periodicals, Inc.
Urine-marking by Canis lupus never occurred when a cache was stocked, rarely occurred during later investigation if some food was still present, but usually occurred soon after the cache was emptied. The animal marking an empty cache was often not the one which had exploited it. Once an empty cache was marked it received little further attention, as opposed to caches that were empty but not urine-marked. Urine-marking may thus enhance foraging efficiency in wolves by signalling that a site contains no more edible food despite the presence of lingering food odors.-from Author
During feeding observations, Canis latrans frequently urine marked the food pile and individual food items that had been carried and dropped some distance from the pile. Food items which had been stolen were often also marked. However, marking of food did not reserve it for the marking animal. Others usually ignored the urine mark and ate the item. The significant increase in urine marking of food during the breeding season suggested that it was involved in the expression of dominance in intrasexual rivalries. Urine marking never occurred when food was cached however, and rarely occurred while the cache still contained food. Once the cache was emptied, urine marking usually occurred: urine marking serves a 'book-keeping' role, indicating that the caches are no longer worth investigating although food odors might still linger.-from Author
Significance
Animals eject fluids for waste elimination, communication, and defense from predators. These diverse systems all rely on the fundamental principles of fluid mechanics, which we use to predict urination duration across a wide range of mammals. In this study, we report a mathematical model that clarifies misconceptions in urology and unifies the results from 41 independent urological and anatomical studies. The theoretical framework presented may be extended to study fluid ejection from animals, a universal phenomenon that has received little attention.
Understanding the role of urine marking in the territorial systems of wild mammals can be difficult, especially for nocturnal cryptic species. Even for common species, such as the red fox Vulpes vulpes, a comprehensive analysis of seasonal and sex differences has not been carried out. Using 6 years of infra-red video monitoring, we compared marking rates between months and between sexes. Urine marking was significantly lower during summer (June–August). Males urine marked significantly more frequently than females during late summer and autumn, but not winter. Males marked more frequently than females also during March. There was no increase during the breeding season for either sex. Our results correlate with previous partial data but demonstrate how urine marking rates vary across the year. They also further support the greater role of males in fox territorial maintenance. Urine marking is lowest during summer when territorial intrusions are least, whilst the higher male urine marking rate in March reflects the period when females are denning. Overall, our results provide the first comprehensive analysis of red fox urine marking rates, contributing to a greater understanding of territoriality and olfactory communication.
The relationship between urine-marking and caching was studied in two captive groups of wolves (Canis lupus). It was found that urine-marking never occurred when a cache was stocked, rarely occurred during later investigations if some food was still present, but usually occurred soon after the cache was emptied. The animal marking an empty cache was often not the one which had exploited it. Once an empty cache was marked it received little further attention, as opposed to caches that were empty but not urine-marked. These results suggest that urine-marking may enhance foraging efficiency in wolves by signalling that a site contains no more edible food despite the presence of lingering food odors.
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.
We analyzed the leadership behavior of breeding and nonbreeding gray wolves (Canis lupus) in three packs during winter in 1997-1999. Scent-marking, frontal leadership (time and frequency in the lead while traveling), initiation of activity, and nonfrontal leadership were recorded during 499 h of ground-based observations in Yellowstone National Park. All observed scent-marking (N=158) was done by breeding wolves, primarily dominant individuals. Dominant breeding pairs provided most leadership, consistent with a trend in social mammals for leadership to correlate with dominance. Dominant breeding wolves led traveling packs during 64% of recorded behavior bouts (N=591) and 71% of observed travel time (N=64 h). During travel, breeding males and females led packs approximately equally, which probably reflects high parental investment by both breeding male and female wolves. Newly initiated behaviors (N=104) were prompted almost 3 times more often by dominant breeders (70%) than by nonbreeders (25%). Dominant breeding females initiated pack activities almost 4 times more often than subordinate breeding females (30 vs. 8 times). Although one subordinate breeding female led more often than individual nonbreeders in one pack in one season, more commonly this was not the case. In 12 cases breeding wolves exhibited nonfrontal leadership. Among subordinate wolves, leadership behavior was observed in subordinate breeding females and other individuals just prior to their dispersal from natal packs. Subordinate wolves were more often found leading packs that were large and contained many subordinate adults.
Carnivores use various scent-marking methods. Observations on the scent marking by urination of 16 free-ranging dogs from two neighbouring groups were recorded in the town of Katwa, West Bengal, India. The frequency of urine marking was higher in males than in females. The seasonal mean (±S.D.) number of markings for individual males varied with a minimum of 4.0 (±1.5) in summer, and a maximum of 31.5 (±9.1) in late monsoon. Similarly, the seasonal mean (±S.D.) number of markings for individual females varied with a minimum of 1.1 (±0.9) in summer, and a maximum of 5.9 (±2.5) in late monsoon. Therefore, the frequency of marking was not the same in every season. The incidence of urine marking was higher in the courting place as well as in the late monsoon. Occasionally, ‘possessive urine marking’ was observed among the alpha males. Urine marking seemed to be linked with scavenging behaviour. The incidence of urine marking mostly by the males near the territorial boundary during trespassing by neighbouring dog(s) showed the evidence of territorial defense. Perhaps to protect the pups, the females marked with a higher rate at the nest site. The dogs, especially the males, marked on strange objects/vehicles perhaps to familiarise the strange objects.Every male and every female performed High leg raising (HLR) and squat postures, respectively. Raised leg display (RLD) was performed only by the males and was influenced by the presence of other dogs. The males showed the RLD mostly in the courting place and also near the territory, perhaps to indicate their dominance, aggressiveness and to threaten the others.
During staged encounters, bush dogs of the same sex showed a high level of aggressive-defensive behaviors and a higher than normal frequency of urine-marking. Where dominance was established during encounters, dominant individuals marked more than subordinates. Where dominance was not clearly established, both participants remained aggressive and showed higher than normal levels of marking. Urine-marking appears to communicate aggressiveness and may be used to compensate for the otherwise relatively inconspicuous aggressive and dominance displays of this species.
Little is known about factors influencing dyadic interactions between dogs in public places. This paper reports on the effect of dog age, gender and size, human gender and the use of a leash on the occurrence of body sniffing, scent-marking, playing games, showing a threat and biting in canine dyads on walks with their owners. Observations of 1870 interacting dogs were made in public places where owners frequently walked their dogs. Dogs off a leash sniffed one another more often than dogs on a leash (P
Urine marking of food and caches was studied in a group of captive coyotes, Canis latrans. During feeding observations, coyotes frequently urine marked the food pile and individual food items that had been carried and dropped some distance from the pile. In addition, food items which had been stolen were often marked. However, marking of food did not reserve it for the marking animal. Others usually ignored the urine mark and ate the item. The significant increase in urine marking of food during the breeding season suggested that it was involved in the expression of dominance in intrasexual rivalries. Cache marking was quite different. Urine marking never occurred when food was cached and rarely occurred while the cache still contained food. However, once the cache was emptied, urine marking usually occurred. Thus at caches, urine marking evidently serves a "book-keeping" role, indicating that the caches are no longer worth investigating although food odors might still linger.
handIed3 times for 10 min. In the test, handIeddogs encountered 2 persons: the handler in the role of the "owner" (OW) and an unfamiliar person (UP), whereasthe 20 nonhandled dogs encountered unfamiliar persons in both roles.Dogs in the handled gr oup exhibitedmore contact seeking with the enteringOW, less physical contact with the UP, less frequent following of the leaving UP, and less standing by the door in the presence of the OW. The specific response of the handled dogs toward the handlerfulfilled the operational criteria of attachment. In shelter conditions, the remarkable demandfor social contact with humans may result in rather fast formingof attachment even in aduit dogs. Ön the basis of human behavioral observations, Bowlby (1958) described social attachment as an asymmetrical social relationship that presumes the dependency of the attached individual on the object of attachment, who can be used as a secure base. The adaptational significance of social attachment may be supplying offspring with resources for survival and with defense agarost predators by ensuring that offspring remain in the vicinity of the parent (Bowlby, 1969). Attachment is claimed to be the basic organizational factor for any species's social structure leading to group formabon. The dependency of the attached individual man- ifests itself in behavioral preferences indicated by special behavior patterns in choice situations (Wickler, 1976). Up to now, only a few researchers have investigated in exper- imental studies the animal-to-human attachment. Most of them described attachment as the result of imprinting-like processes in a sensitive period, which manifest in proximity seeking and prox-
In female canids, including domestic dogs (Canis familiaris), scent-marking with urine is thought to convey information on reproductive state, yet little is known about how urinary behaviour changes across specific stages of the oestrous cycle. We measured urinary behaviour (proportion of directed urinations) of 10 intact Jack Russell Terriers across anoestrus, proestrus, and oestrus during walks in familiar and novel environments. Females ranged in age from 1.3 to 8.7 years. We assessed stage of oestrus using vaginal cytology, behaviour, and physical signs. Proportions of directed urinations were higher during proestrus and oestrus than anoestrus, and were higher in older females than younger females. Our findings indicate that in female Jack Russell Terriers, scent-marking with urine advertises reproductive state and continues to develop in adulthood. Additional data are needed to determine if these findings generalize to female dogs of other breeds and mixed breeds.
Every year sees an increase in the number of dogs admitted to rescue shelters. However well these dogs are cared for in the shelter it cannot be ignored that being in such a situation is stressful and the time spent in the shelter may change the dogs' behaviour which may in turn influence their chances of being bought from the shelter. This research examined the behaviour of stray and unwanted dogs on their first, third and fifth days in an Ulster Society for the Prevention of Cruelty to Animals (USPCA) shelter. A questionnaire was also distributed to members of the public to determine how popular the USPCA was as a place from where to purchase a dog, and what factors about a dog's physical characteristics, behaviour and environment influenced potential buyers. Results revealed no significant difference between the behaviour of stray and unwanted dogs although the public viewed stray dogs as much less desirable than unwanted dogs. Time in the shelter had no adverse effects on the dogs' behaviour. Indeed those changes which did occur during captivity, dogs being more relaxed in the presence of people and eating food more quickly, may be considered as positive changes. The USPCA was viewed as a popular place from which to buy a dog. Off actors influencing the public's choice, the dog's environment and behaviour appeared more important than its physical characteristics. The presence of a toy in the dog's cage greatly increased the public's preference for the dog, although the toy was ignored by the dog. The welfare implications of sheltering dogs are discussed
In many species, body size is a key determinant of the outcome of agonistic interactions, and receivers are expected to attend to size cues when assessing competitors' signals. Several mammal vocalizations, including domestic dog growls, encode reliable information about caller body size in the dispersion of formant frequencies. To test whether adult domestic dogs attend to formant dispersion when presented with the growls of their conspecifics, we played recordings of resynthesized growls where the size-related variation in formant frequency spacing was manipulated independently of all other parameters. Subjects from three different size groups (small, medium and large dogs) were presented with playbacks of growls where formant frequencies had been rescaled to correspond to a dog 30% smaller or 30% larger than themselves. While large dogs systematically displayed more motivation to interact when growls simulated a smaller intruder, small dogs did not respond differentially to the playback conditions. However, the small dogs responded significantly less than all other size groups to both playback conditions. Our results suggest that domestic dogs are able to perceive size-related information in growls, and more specifically that they are able to adapt their behavioural response as a function of the perceived intruder's size relative to their own.
The tail of dogs and allies (Canidae) is important for intraspecific communication. We used a life-sized dog model and varied the tail length and motion as an experimental method of examining effects of tail-docking on intraspecific signaling in domestic dogs, Canis famil-iaris. We videotaped interactions of 492 off-leash dogs and quantified size and behaviour of approaching dogs to the model's four tail conditions (short/still, short/wagging, long/still, long/wagging). Larger dogs were less cautious and more likely to approach a long/wagging tail rather than a long/still tail, but did not differ in their approach to a short/still and a short/wagging tail. Using discriminant analyses of behavioural variables, dogs responded with an elevated head and tail to a long/wagging tail model relative to the long/still tail model, but did not show any differences in response to tail motion when the model's tail was short. Our study provides evidence that a longer tail is more effective at conveying different in-traspecific cues, such as those provided by tail motion, than a shorter tail and demonstrates the usefulness of robotic models when investigating complex behavioural interactions.
The urine-marking behaviour of captive, paired male and female bush dogs, Speothos venaticus, was investigated during pair formation and maintenance. Urine-marking frequencies decreased as pair members became familiar, but urine-marking remained an important aspect of the bush dog's precopulatory display. Paired females exhibited a handstand posture, as well as several other urine-marking behaviours less often or never shown by males. The onset of urine-marking occurred later in female versus male juveniles living in the family group. Behavioural responses suggest that bush dogs can discriminate between male and female urine and that female handstanding may serve as a visual signal in sexual recognition. Sexual differences in urine-marking behaviours are interpreted in relation to intrasexual competition for the high level of parental care exhibited by males in this monogamous species.
This paper reviews experimental and field studies on scent marking behaviour. The occurrence and effects of scent marking are considered in particular, and a number of areas for further research are made apparent. Marking behaviour in mammals is often stated to be ‘territorial’ or, more specifically, to play a role in territorial defence. In fact there is a shortage of evidence to support this view; many of the relevant observations are anecdotal or interpreted with preconceived notions of function in mind. While marking is clearly associated with aggressive behaviour in many species and may therefore be related in some way to territorial behaviour, its role in aggression is not understood. Moreover, there is evidence to support a number of other theories of function some of which are unrelated to territory. It seems that, as with any other mode of communication, scent marking has become adapted for use in a variety of contexts. It probably has more than one function in any one species and different functions in different species.
Scent-marking was studied in wolves (Canis lupus) along 133 km of tracks in northern Minnesota during winters of 1975 to 1976 and 1976 to 1977 and in two captive packs and four captive pairs for various periods. Lone wolves, which possess neither mates nor territories, rarely marked by raised-leg urination and defaecated and urinated less along roads and trails, where territorial pairs and packs generally marked. Newly formed pairs marked the most, eventually decreasing their rates to those of established packs. Generally, wolves that scent-marked also bred, whereas non-marking wolves usually did not breed. Scent-marking apparently is important to the success of courtship in new pairs and to reproductive synchrony in established pairs, as well as serving a territorial function.
The effects of testicular and ovarian hormones on urinary frequency and posture were studied in 8 groups of dogs. Three of the 4 female groups had been exposed to androgenic stimulation in utero, neonatally, or both in utero and in infancy. The fourth group of females consisted of ovariectomized controls. Males were intact or castrated as adults, as juveniles, or as neonates. Frequency of urination was increased in all groups by estrogen and by testosterone. Urinary posture was unaffected in males by castration in the adult or juvenile stages. Neonatal castration of males resulted in periodic regression from the adult male posture to the immature male posture. Control and prenatally androgenized females urinated in the feminine position. Females treated with large amounts of testosterone in infancy showed a limited degree of masculinization of urinary posture. Females exposed to testosterone before and immediately after birth urinated as females about 50% of the time and as adult males for approximately half of their urinations. Exogenous estrogen or androgen administered in adulthood had no effect on posture assumed for urination.
Development of urinary behavior from birth to adulthood was observed in six groups of beagles including normal males (NM), normal females (NF), males castrated soon after birth (CM), males castrated soon after birth and treated with testosterone (T) for the next 90 days (CMT), females exposed to T in utero (FTU), and females exposed to T in utero and during infancy, i.e., the first 30-40 days postpartum (FTUI). Prenatal treatment with T had masculinizing effects on juvenile urinary behavior in FTU and FTUI. On the other hand normal development of fully adult masculine urinary patterns in males and females necessitated both prenatal and postnatal androgenic stimulation. It was not necessary that T be present at the time the overt behavior developed. For example, adult male behavior appeared in FTUI at the same time as in NM, i.e., 6-10 months, although the supply of exogenous androgen in FTUI had been exhausted within 30-40 days after birth. CMT showed precocious development of all components of the adult male pattern. Development of adult responses was markedly retarded in most CM, and their performance did not equal that of NM at 23 months. They were then injected with TP which promptly evoked completely normal male urinary behavior. It is tentatively concluded that T acting before birth and during the juvenile period "prepares" critical CNS mechanisms so that when general maturation reaches the appropriate point adult male behavior develops. Although the preparatory role of T is essential, the behavior is not dependent on T after it has developed.
Twelve distinct elimination postures were recorded during the present study. Females employed a wider variety of urination postures, eight compared to four in the males. On the other hand, more males urinated during more tests and did so more frequently than the females. Too, the males showed a considerably lower urination latency than the bitches. Females tended to evacuate the bladder all at once while the males would deposit a small amount of urine many times and continue to pseudo-urinate even though no urine was expelled. In the males, urination appeared to be oriented toward the scent of other males or toward conspicuous vertical objects while female urinations tended to be distributed at random. Although most elimination by the females was aimed at the ground, a substantial number of their urinations were directed toward vertical targets as well. The defecation rate was low in both sexes, being slightly higher in the males. Five defecation postures were used by the males as opposed to two by the females. Of the postures observed, two were formerly thought to be used for urination only. Both sexes showed some defecation on vertical targets. Almost one-fourth of male defecations were placed on the fence. A small amount of ground scratching was done, most of it by the males, who scratched only after urination. Scratching appears to function as part of the marking behavior of the dog. It may help to disperse an animal's odor over a wide area as well as making a conspicuous mark on some surfaces.
When adult male dogs were placed in pens previously occupied by other dogs, a mean of 24.6 urine marks were deposited in the first 2 hrs; marking frequency was then reduced to about two to five marks per 2 hr over the remainder of the day and on subsequent days. A high rate of marking behavior was again observed when subjects were placed in a special marking pen for 5 min. These observations demonstrate some similarities between urine marking in male dogs and scent marking in males of some rodent species. Following castration no quantitative changes in urine marking in the special marking pen were evident after 5 mo of postcastration testing even though a decline in mating behavior occurred within 2 mo. This contrasts with the rather prompt reduction in scent marking in male rodents following castration.
We observed 49 coyotes, Canis latransfrom five resident packs for 2456 h and five transient coyotes for 51 h from January 1991 to June 1993 in the Lamar River Valley, Yellowstone National Park, Wyoming, U.S.A. During these observations we recorded 3042 urinations, 451 defecations, 446 ground scratches and 743 double-marks. The rate of scent-marking (via urination) was influenced by the social organization (resident versus transient) to which the coyote belonged, the social class (alpha, beta or pup) of the animal and the time of the year. Transient coyotes scent-marked at a lower rate than did members of a resident pack. Within the resident packs, alpha coyotes scent-marked at a higher rate than beta coyotes (adults and yearlings subordinant to alphas, but dominant over pups) and pups. Alpha coyotes increased their rate of marking during the breeding season; beta and pup coyotes performed scent-marks at a relatively constant rate throughout the year. There was no influence of social class or time of year on the rate of defecation. The rate of double-marking was highest among alpha coyotes with a peak during the breeding season. Alpha coyotes ground-scratched at a higher rate than did beta and pup coyotes. Alpha and beta coyotes scent-marked more than expected along the periphery of the territory compared to the interior; pups marked in the interior and edge in proportion to expected frequencies. Double-marking and ground-scratching were higher than expected along the periphery of the territory. The distribution of defecations was not different from expected along the edge versus the interior of the territory. Pack size did not influence the rate of scent-marking performed by individuals in the pack or by the alpha pair. We concluded that alpha coyotes were the primary members of the resident pack involved in scent-marking. The large coyote packs and the high rate of marking by the alpha pairs were parallel to the scent-marking behaviour displayed by wolves, C. lupusto a greater extent than previously reported. Scent-marks appear to provide internal information to the members of the resident pack (internal map of territory, breeding condition, reproductive synchrony) and enhance demarcation of territorial boundaries.Copyright 1997 The Association for the Study of Animal Behaviour1997The Association for the Study of Animal Behaviour
Many mammals use scent marks to advertise territory ownership, but only recently have we started to understand the complexity of these scent signals and the types of information that they convey. Whilst attention has generally focused on volatile odorants as the main information molecules in scents, studies of the house mouse have now defined a role for a family of proteins termed major urinary proteins (MUPs) which are, of course, involatile. MUPs bind male signalling volatiles and control their release from scent marks. These proteins are also highly polymorphic and the pattern of polymorphic variants provides a stable ownership signal that communicates genome-derived information on the individual identity of the scent owner. Here we review the interaction between the chemical basis of mouse scents and the dynamics of their competitive scent marking behaviour, demonstrating how it is possible to provide reliable signals of the competitive ability and identity of individual males.
Urinary cortisol levels (based on cortisol : creatinine ratios) were evaluated in a randomly selected sample of shelter dogs kennelled over a 31-day period. Urine was collected on days 2, 5, 10, 17, 24 and 31 (with day 1 referring to the day of admittance to the shelter). Cortisol levels peaked on day 17 and steadily declined thereafter, although a high degree of individual variation was found, with cortisol levels peaking sooner in some dogs. Cortisol levels in kennelled dogs were significantly higher on all days except d 31 than the baseline measures taken from 20 dogs in their home environments. There were no differences between cortisol levels in male and female dogs on each day of sampling and there was no significant linear correlation between age and cortisol levels. The results are discussed in relation to stress management and the welfare of kennelled dogs.
this paper we have argued that the study of the proximate mechanisms underlying vocal behavior, both physiological and cognitive, is a necessary part of the study of the evolution of communication, and in particular for analyzing honesty in communication. In the first part we surveyed basic principles of vocal production in terrestrial vertebrates, and the morphological diversity of their production systems. We then provided some examples of the interactions between acoustics and anatomy that can enforce honesty, or subvert it. In the second part we examined the evidence for cognitive mechanisms that allow animals to produce deceptive calls, as well as "retaliatory" perceptual mechanisms that allow perceivers to accurately identify and ignore (and in some cases even punish) the deceivers. Both vocal production mechanisms, and cognitive mechanisms controlling vocalization, play a crucial role in determining what is possible or impossible in a particular species' communication system. A better understanding of these mechanisms can lead to rich insights into the evolution of acoustic communication. Fitch & Hauser p. 38 For the reader already interested in mechanism, the chapter also provided illustrations of the value of an ultimate evolutionary viewpoint. An evolutionary perspective proves valuable both for identifying functional problems that are solved by communicators, and for using phylogenies and the comparative method as tools to identify and understand widespread selective pressures and functional constraints. The species we observe today are the outcome of a long dynamic process of coevolution and interaction. Signalers' ability to avoid, repel or attract predators, competitors and potential mates has played a critical role in the evolution of their acoustic si...
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