Article

Urine marking by free-ranging dogs ( Canis familiaris) in relation to sex, season, place and posture

Authors:
  • Katwa Bharati Bhaban
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Abstract

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.

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... Scent marking has been studied in detail in several members of Canidae. Possible functions range from defending a territory (freeranging dogs: Cafazzo et al., 2012;Pal, 2003;coyotes: Gese and Ruff, 1997; gray wolves: Peters and Mech, 1975) to providing olfactory and possibly visual landmarks, which aid in orientation and making objects within territories more familiar (free-ranging dogs: Cafazzo et al., 2012;Pal, 2003;coyotes: Gese and Ruff, 1997) to indicating characteristics of food, such as location, ownership, or that a cache is empty (free-ranging dogs: Cafazzo et al., 2012;Pal, 2003;coyotes: Harrington, 1982;gray wolves: Harrington, 1981;red foxes: Henry, 1977). With respect to social interactions, scent marking may establish or reinforce social status (companion dogs: Lisberg and Snowdon, 2011;free-ranging dogs: Cafazzo et al., 2012;coyotes: Gese and Ruff, 1997;gray wolves: Peterson et al., 2002;bush dogs (Speothos venaticus): Biben, 1982) and bonds between members of a breeding pair (coyotes: Gese and Ruff, 1997;gray wolves: Rothman and Mech, 1979;bush dogs: Porton, 1983). ...
... Scent marking has been studied in detail in several members of Canidae. Possible functions range from defending a territory (freeranging dogs: Cafazzo et al., 2012;Pal, 2003;coyotes: Gese and Ruff, 1997; gray wolves: Peters and Mech, 1975) to providing olfactory and possibly visual landmarks, which aid in orientation and making objects within territories more familiar (free-ranging dogs: Cafazzo et al., 2012;Pal, 2003;coyotes: Gese and Ruff, 1997) to indicating characteristics of food, such as location, ownership, or that a cache is empty (free-ranging dogs: Cafazzo et al., 2012;Pal, 2003;coyotes: Harrington, 1982;gray wolves: Harrington, 1981;red foxes: Henry, 1977). With respect to social interactions, scent marking may establish or reinforce social status (companion dogs: Lisberg and Snowdon, 2011;free-ranging dogs: Cafazzo et al., 2012;coyotes: Gese and Ruff, 1997;gray wolves: Peterson et al., 2002;bush dogs (Speothos venaticus): Biben, 1982) and bonds between members of a breeding pair (coyotes: Gese and Ruff, 1997;gray wolves: Rothman and Mech, 1979;bush dogs: Porton, 1983). ...
... Scent marking has been studied in detail in several members of Canidae. Possible functions range from defending a territory (freeranging dogs: Cafazzo et al., 2012;Pal, 2003;coyotes: Gese and Ruff, 1997; gray wolves: Peters and Mech, 1975) to providing olfactory and possibly visual landmarks, which aid in orientation and making objects within territories more familiar (free-ranging dogs: Cafazzo et al., 2012;Pal, 2003;coyotes: Gese and Ruff, 1997) to indicating characteristics of food, such as location, ownership, or that a cache is empty (free-ranging dogs: Cafazzo et al., 2012;Pal, 2003;coyotes: Harrington, 1982;gray wolves: Harrington, 1981;red foxes: Henry, 1977). With respect to social interactions, scent marking may establish or reinforce social status (companion dogs: Lisberg and Snowdon, 2011;free-ranging dogs: Cafazzo et al., 2012;coyotes: Gese and Ruff, 1997;gray wolves: Peterson et al., 2002;bush dogs (Speothos venaticus): Biben, 1982) and bonds between members of a breeding pair (coyotes: Gese and Ruff, 1997;gray wolves: Rothman and Mech, 1979;bush dogs: Porton, 1983). ...
... Males urinate and countermark (mark on or near existing scent marks) more frequently than do females and direct more of their urinations at targets in the environment; typically, adult males raise a hindlimb to urinate whereas adult females squat (Beach, 1974;Bekoff, 1979a;Lisberg and Snowdon, 2011;Martins and Valle, 1948;Sprague and Anisko, 1973). Sexual dimorphism in urine-marking behavior has been reported for freeranging dogs (Cafazzo et al., 2012;Pal, 2003), as well as for dogs maintained in either laboratory colonies or homes (Beach, 1974;Lisberg and Snowdon, 2011;Martins and Valle, 1948;Sprague and Anisko, 1973). In contrast to urinary behavior, sex differences do not characterize either defecation (Sprague and Anisko, 1973) or ground scratching (backward scraping of the ground with the front feet, hind feet, or both performed by some dogs after urination or defecation; Bekoff, 1979a). ...
... At least nine functions have been suggested for scent marking in members of Canidae (Spotte, 2012). The six with the most empirical support include: 1) defending a territory (free-ranging dogs: Cafazzo et al., 2012;Pal, 2003;coyotes: Gese and Ruff, 1997;gray wolves: Peters and Mech, 1975); 2) establishing or reinforcing social status (companion dogs: Lisberg and Snowdon, 2011;free-ranging dogs: Cafazzo et al., 2012;coyotes: Gese and Ruff, 1997;gray wolves: Peterson et al., 2002;bush dogs (Speothos venaticus): Biben, 1982); 3) indicating female sexual status (companion dogs: Wirant et al., 2007; free-ranging dogs: Cafazzo et al., 2012;Pal, 2003); 4) establishing and maintaining pair bonds (coyotes: Gese and Ruff, 1997; gray wolves: Rothman and Mech, 1979;bush dogs: Porton, 1983); 5) providing olfactory landmarks within territories to either assist in orientation or make strange objects or areas more familiar (free-ranging dogs: Cafazzo et al., 2012;Pal, 2003;coyotes: Gese and Ruff, 1997); and 6) indicating either the location of food, a claim to food, or an empty food cache (free-ranging dogs: Cafazzo et al., 2012;Pal, 2003;coyotes: Harrington, 1982;gray wolves: Harrington, 1981; red foxes: Henry, 1977). Evidence to support these functions typically includes comparative data on rates of urine marking by individuals either of different sex, age class, or social status, or in different contexts (e.g., at the boundaries versus the interior of a territory). ...
... At least nine functions have been suggested for scent marking in members of Canidae (Spotte, 2012). The six with the most empirical support include: 1) defending a territory (free-ranging dogs: Cafazzo et al., 2012;Pal, 2003;coyotes: Gese and Ruff, 1997;gray wolves: Peters and Mech, 1975); 2) establishing or reinforcing social status (companion dogs: Lisberg and Snowdon, 2011;free-ranging dogs: Cafazzo et al., 2012;coyotes: Gese and Ruff, 1997;gray wolves: Peterson et al., 2002;bush dogs (Speothos venaticus): Biben, 1982); 3) indicating female sexual status (companion dogs: Wirant et al., 2007; free-ranging dogs: Cafazzo et al., 2012;Pal, 2003); 4) establishing and maintaining pair bonds (coyotes: Gese and Ruff, 1997; gray wolves: Rothman and Mech, 1979;bush dogs: Porton, 1983); 5) providing olfactory landmarks within territories to either assist in orientation or make strange objects or areas more familiar (free-ranging dogs: Cafazzo et al., 2012;Pal, 2003;coyotes: Gese and Ruff, 1997); and 6) indicating either the location of food, a claim to food, or an empty food cache (free-ranging dogs: Cafazzo et al., 2012;Pal, 2003;coyotes: Harrington, 1982;gray wolves: Harrington, 1981; red foxes: Henry, 1977). Evidence to support these functions typically includes comparative data on rates of urine marking by individuals either of different sex, age class, or social status, or in different contexts (e.g., at the boundaries versus the interior of a territory). ...
Article
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.
... Olfactory cues are used for territory marking, individual recognition and as a method for leaving long term messages (3). Marks may be made on novel or familiar objects, territory, or scent marks of other individuals, or the animal may mark itself (6,33,51,52). Unfamiliar urine, faeces or objects may be marked simply to reduce novelty (3). Feeding places can also be marked, probably increasing the efficiency of food source utilisation (6,52). ...
... Unfamiliar urine, faeces or objects may be marked simply to reduce novelty (3). Feeding places can also be marked, probably increasing the efficiency of food source utilisation (6,52). ...
... In feral dogs females marked more frequently near the nest side after litter production and when they were in oestrus (52). One of the messages that urine contains is about female reproductive status and oestrus females may urinate more, leaving long-distance messages for males. ...
... year-round territorial defence mainly by males (Gese & Ruff 1997;Pal 2003;Núñez & De Miguel 2004;Arnold et al. 2011;Cafazzo, Natoli & Valsecchi 2012) and male foxes guard their mates during the short oestrus period (Baker et al. 2004a;Kitchen et al. 2005a;Ralls et al. 2013), which may limit foraging opportunities (Alberts et al. 1996;Girard-Buttoz et al. 2014). In foxes, changes in group composition are promoted by population density and are accompanied by changes in mating system (Iossa et al. 2009), probably when males lose their ability to monopolise reproduction (Say et al. 1999;Wright et al. 2010). ...
... This greatly increases mate competition between males (and females) and the reduction in male foraging effort I observed in winter contributes to their significant loss of body mass between winter and spring (Saunders et al. 1993). Along with increased extraterritorial movement (this study; Soulsbury et al. 2011), this reflects the effort male foxes exert to secure copulations, a motivation common to all male mammals, as reflected by sex differences in scent marking behaviour, i.e. males mark more and at the territory boundary, while females mark resource patches (Pal 2003;Parker 2010;Clapham et al. 2014). In fitting with this, I observed greater foraging efficiency by females than males. ...
Thesis
Full-text available
The red fox (Vulpes vulpes) is a contentious species of global importance as a predator, competitor, vector of disease and pest. Understanding their social system is essential for successful species management and mitigating human-wildlife conflict. Foxes are solitary foragers that form groups in certain circumstances. It is unclear whether these groups are beneficial or simply the ‘best of a bad job’ due to ecological constraints. Further, little is known about how stable or cohesive fox groups are, which can have implications for disturbance or removal. I addressed these issues to further our understanding of how fox groups operate. I set up camera traps in residential gardens where foxes were fed (food patches), to study social and competitive relationships in the high density fox population of suburban Bristol, UK. I collected over 152,000 photos of foxes and identified 192 individuals. Social groups were difficult to define, but the most reliable definition encompassed shared space use, a sighting threshold and the number of social connections. Group membership was relatively stable; foxes associated in communities, mainly within their territory boundary and maintained long term relationships that lasted until death or emigration. However, in all seasons the majority of relationships lasted less than a day and were probably between foxes from different social groups, indicating that intergroup contacts were not uncommon and occurred year round. Food patches were hotspots for sociality. Foxes improved their foraging efficiency by selecting high quality patches and coinciding their foraging activity with anticipated food availability, which increased contact rates at high quality patches. Females foraged according to their seasonal energetic demands, while males reduced their foraging effort in the winter mating season in favour of mate-searching behaviour. This contributed to a significant alteration in social structure in winter, with an increased rate of territory intrusion by strangers, a greater proportion of short term relationships and reduced social connectivity, demonstrating a role of females in maintaining group cohesion in winter. Dominant foxes occupied central network positions and therefore had a major influence on group connectedness; the demise of a dominant male led to significant social perturbation in one territory, supporting the importance of breeders in canid groups. Dominance also facilitated priority access to food, but so too did resident group membership, perhaps through familiarity with local resources and conspecifics. Subordinates compensated for intragroup competition by utilising lower quality patches and risky extraterritorial foraging, which was observed at an unexpectedly high rate year round. Contrary to the resource dispersion hypothesis, this indicated that group size was not limited by within-territory resources. To my knowledge, this is the most detailed study of fox sociality to date. Despite low contact rates fox groups were relatively stable and more similar in complexity to other canids than previously acknowledged. High individual flexibility in space use and year-round extraterritorial movement may have functioned to mitigate competition, but also explained how foxes responded so rapidly to local demographic change, providing evidence for the futility of management by removal.
... At least one aspect of canine scent-marking behavior is sensitive to fearful or stressful conditions: adult male dogs that used the raised-leg urinary posture typical of mature males temporarily reverted in fearful situations to using the juvenile lean-forward posture in which all four feet remain on the ground [33,34]. Consistent with this finding, we reported that the percent of urinations in which adult male dogs used the raised-leg posture was lower in our study shelter (73%; [35]) than reported for mature male dogs living under other conditions (94%-97%; [36][37][38][39]). We found a similar effect for female dogs: 6% of urinations by adult females involved raising a hindlimb at our study shelter [35] compared with 19%-37% for adult female dogs living under other conditions [36][37][38][39]. ...
... Consistent with this finding, we reported that the percent of urinations in which adult male dogs used the raised-leg posture was lower in our study shelter (73%; [35]) than reported for mature male dogs living under other conditions (94%-97%; [36][37][38][39]). We found a similar effect for female dogs: 6% of urinations by adult females involved raising a hindlimb at our study shelter [35] compared with 19%-37% for adult female dogs living under other conditions [36][37][38][39]. These observations suggest that monitoring scent-marking behavior of dogs during walks might be a useful way to assess how shelter dogs respond to the sex of an unfamiliar walker. ...
Article
Full-text available
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.
... Concerning the free-roaming pet dogs, we can assume that they were motivated to roam not only for food, but also for sociality [12,20,48,49,109]. Cachupín could only have limited interactions with leashed pet dogs and was actually seen with stray dogs. ...
... Here, efficient survival strategies are at play, meaning that the animal will respond to any external threat in the form of attack, submission or flight. For canids, a territory/niche is a living space (defined as an island by Boitani [17]), marked by odor (urine), where they can perform the actions needed for their survival [48,49,123], namely finding food and water, resting in a safe place, mating and rearing offspring. Each territory is home to a number of individuals that varies not just according to the species, but as a function of other factors too [46]. ...
Article
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Stray dogs are a common sight in cities all over the world, especially in Latin America, but research on their behavior is scarce. Studying their very considerable presence in Concepción (Chile) provided a unique opportunity to learn more about the extent of the sociality and territoriality patterns of the dog species. Interestingly, a wide range of socialities with humans (and with other dogs) were shown to be dependent on human activities and urban zoning signaled by passages, physical boundaries and landmarks. New forms of sociality were also evidenced, with dogs exhibiting intermediate degrees of sociality between the pet and stray dog categories. We postulate that this unique diversity of sociospatial positioning and level of adjustment (e.g., dogs using crosswalks either alone or with people) is made possible by the city’s specific human culture and range of urban areas. The dog species thus exhibits a considerable potential for social and spatial adjustment. The fact that it depends on the spatial layout and human culture of their environment explains the presence of dogs wherever humans are. Furthermore, it has implications for coping with the presence of dogs in numerous and varied human societies.
... However, the signal decay due to precipitation, ultraviolet radiation and bacterial decomposition has rarely been investigated empirically 20,21 . For example, free-ranging dogs have been observed to scent mark more during the wet season 22 , thus increasing the cost of territory defence at these times. It is possible that changes in rainfall or ultraviolet radiation, due to climate change for instance 23 , may have an impact on the economics of territory defence for scent marking species more generally 10 . ...
... www.nature.com/scientificreports/ The rate of scent marking at the territory boundary may provide insight into the territorial pressure exerted by conspecifics 22 . The spatial positioning of scent marks within the territory may also indicate their function. ...
Article
Full-text available
For canid species, scent marking plays a critical role in territoriality, social dynamics, and reproduction. However, due in part to human dependence on vision as our primary sensory modality, research on olfactory communication is hampered by a lack of tractable methods. In this study, we leverage a powerful biologging approach, using accelerometers in concert with GPS loggers to monitor and describe scent-marking events in time and space. We performed a validation experiment with domestic dogs, monitoring them by video concurrently with the novel biologging approach. We attached an accelerometer to the pelvis of 31 dogs (19 males and 12 females), detecting raised-leg and squat posture urinations by monitoring the change in device orientation. We then deployed this technique to describe the scent marking activity of 3 guardian dogs as they defend livestock from coyote depredation in California, providing an example use-case for the technique. During validation, the algorithm correctly classified 92% of accelerometer readings. High performance was partly due to the conspicuous signatures of archetypal raised-leg postures in the accelerometer data. Accuracy did not vary with the weight, age, and sex of the dogs, resulting in a method that is broadly applicable across canid species’ morphologies. We also used models trained on each individual to detect scent marking of others to emulate the use of captive surrogates for model training. We observed no relationship between the similarity in body weight between the dog pairs and the overall accuracy of predictions, although models performed best when trained and tested on the same individual. We discuss how existing methods in the field of movement ecology can be extended to use this exciting new data type. This paper represents an important first step in opening new avenues of research by leveraging the power of modern-technologies and machine-learning to this field.
... Communication via scents plays an important role in dogs' reproductive behaviour. Bitches signal their reproductive status through urine marks and vaginal secretions [94], whose odour is extremely attractive for other dogs [95]. It elicits a specific reaction in male dogs, which deposit their own urine on or near to the females' one as a signal for courtship [94]. ...
... Bitches signal their reproductive status through urine marks and vaginal secretions [94], whose odour is extremely attractive for other dogs [95]. It elicits a specific reaction in male dogs, which deposit their own urine on or near to the females' one as a signal for courtship [94]. ...
Article
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Simple Summary Communication takes place between members of the same species, as well as between heterospecific individuals, such as the long co-habitation process and inter-dependent relationship present in domestic dogs and humans. Dogs engage in visual communication by modifying different parts of their body; in tactile communication; and also in auditory and olfactory communication, with vocalizations and body odours, respectively. The aim of this review is to provide an overview of the recent literature about dog communication, describing the different nature of the signals used in conspecific and heterospecific interactions and their communicative meaning. Lateralized dog brain patterns underlying basic neural mechanisms are also discussed, for both conspecific and heterospecific social communication. Abstract Dogs have a vast and flexible repertoire of visual, acoustic, and olfactory signals that allow an expressive and fine tuned conspecific and dog–human communication. Dogs use this behavioural repertoire when communicating with humans, employing the same signals used during conspecific interactions, some of which can acquire and carry a different meaning when directed toward humans. The aim of this review is to provide an overview of the latest progress made in the study of dog communication, describing the different nature of the signals used in conspecific (dog–dog) and heterospecific (dog–human) interactions and their communicative meaning. Finally, behavioural asymmetries that reflect lateralized neural patterns involved in both dog–dog and dog–human social communication are discussed.
... Both male and female dingoes used standing raised leg urination only, with or without ground scratching, when scent marking the star pickets with urine. Canids can use a range of postures when scent marking (Asa et al. 1985;Pal 2003), and females are often recorded using squat urinations (Rothman and Mech 1979;Wells and Bekoff 1981;Sillero-Zubiri and Macdonald 1998). Standing raised leg urination and ground scratching is associated with scent marking by both the male and female of the dominant pair, and often used for demarcation of territorial boundaries (Wells and Bekoff 1981;Sillero-Zubiri and Macdonald 1998;Allen et al. 1999). ...
... However, it seems likely that when working Maremmas scent-mark their territory, dingoes will investigate these marks, and engage in olfactory communication in response. Most species of canid are territorial (Macdonald and Sillero-Zubiri 2004), including domestic dogs (Font 1987;Pal 2003) and dingoes (Thomson 1992;Corbett 2001). Scent marking with scats, urine and ground scratching are used for social communication in all of these species, including advertisement of ownership and territory boundaries (Sillero-Zubiri and Macdonald 1998). ...
Article
The behavioural mechanisms by which livestock guardian dogs (LGDs) protect livestock from wild predators are not yet fully understood. LGD urine could play a part, as scent-marking the boundaries of a territory could signal occupation of the area to predators. Past selection for dogs that were most effective in deterring predators could have resulted in LGDs that produce urine with predator-deterrent properties. In this research, 28 captive dingoes (14 male and 14 female) were tested for their response to urine marks of LGDs (Maremma sheepdogs), herding dogs (Border Collies) and other dingoes, with distilled water used as a control. The response of the dingoes to the scents was measured using eight variables. For most variables, the response to the test scents was not statistically different from the response to the control. Test minus control was calculated for each test scent category, and used to compare responses between different test scents. The response to Maremma urine was similar to the response to Border Collie urine, and resembled a reaction to a conspecific. We found no evidence of predator-repellent properties of LGD urine. Our results suggest that dingoes readily engage in olfactory communication with Maremmas. It therefore seems likely that they would recognise territorial boundaries created by working Maremmas.
... Communication tactics using olfactory methods (including scent-marking) have been widely investigated (Rothman and Mech 1979;Barrette and Messier 1980;Van Heerden 1981;Wells and Bekoff 1981;Asa et al. 1985;Paquet 1991;Gese and Ruff 1997;Sillero-Zubiri and Macdonald 1998;Allen et al. 1999;Bekoff 2001;Briscoe et al. 2002;Peterson et al. 2002;Pal 2003;Wirant and McGuire 2004;Barja et al. 2005;. Scent-marking is the deposition of scented secretions in the environment (Barrette and Messier 1980) and involves the use of semiochemicals, which are any chemical substances produced by an animal that are used in communication. ...
... Scent-marking via 9 urination is easy to distinguish from simple elimination because it involves some distinct behaviour such as directional flow (Kleiman 1966), usually while adopting either a raised-leg or forward-lean posture (Allen et al. 1999). In addition, the urine is aimed at some environmental landmark (primarily vertical objects, often over previous urine marks) , a relatively small amount of urine is expelled, and canids sometimes mark after first investigating and sniffing several spots (Pal 2003). Another behaviour associated with scent-marking is ground-scratching, where, after urination and/or defecation, the canid paws the ground vigorously (Kleiman 1966;Barrette and Messier 1980;Wirant and McGuire 2004). ...
Technical Report
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Canids use olfactory communication to convey a variety of messages, including identification of social status and home range, attracting sexual partners, and information for the internal orientation of members of the resident pack. Another form of olfactory communication in canids is territorial scent-marking, in which one pack or one individual leaves an identifying semiochemical message and acquires information about other packs or individuals in a given region. In recent years, investigation into the use of artificial boundaries derived from predators’ semiochemical signals to manage human/predator conflict has shown significant promise. Synthesised semiochemicals of African wild dogs have been trialled for restriction of their movement and reduction of conflict with local farmers. In Australia, dingoes (Canis lupus dingo) (and hybrids with domestic dogs Canis lupus familiaris) cause substantial economic loss to agriculture and are actively controlled in many parts of their range. However, dingoes are also considered to be protected native species in parts of their range. This contradiction in status requires the development of innovative and non-lethal control tools. This project extends previous investigations on the chemical composition of dingo urine by first collecting and analysing the composition of urine from six dominant male dingoes, and second by undertaking pen trials using two potential semiochemicals to investigate their potential for influencing dingo behaviour. We identified 28 chemicals in dingo urine, comprising 10 functional groups (combinations of chemical molecules that react in similar ways). Although many of the individual chemicals were unique to dingoes, the functional groups found in dingo urine were also found in other predators such as wolves (Canis spp.), coyotes (Canis latrans), domestic dogs (Canis lupus familiaris), red foxes (Vulpes vulpes), and African wild dogs (Lycaon pictus). Several potential semiochemicals that could play a role in marking territory were identified as potential candidates for assessment. However, many were not commercially available. In the pen trials, we exposed 15 dingoes to two semiochemicals and a control (water) over 8 days with a repeated cross-over study design. The results indicated that it is possible to manipulate dingo behaviour using semiochemicals extracted from dominant male dingo urine. However, other factors such as sex, social status and weather also played a role in explaining differences in behaviour. For compounds to be considered candidates for use in managing the spatial behaviour of canids, they need to be species specific, potentially airborne, persistent, and present in urine and not in faeces. It is possible that important chemicals may be found in the very low levels of substance concentration (e.g. pico- (1E−12) to nanomolar (1E−9) gas phase concentrations), which are easily within the range of a dingo’s sense of smell but close to the limits of detection for the analytical tools available. This, along with the range of semiochemicals in urine, highlights the complex nature of identifying which combination of compounds found in urine is responsible for communication of a specific message. Future investigations of dingo semiochemicals should focus on extracting more detail regarding the smaller fractions of the urine profile (Apps et al. 2013) and on trialling combinations of the most promising chemicals. Research into semiochemicals in other species of canids continues, and sharing approaches to selecting semiochemicals should be considered. Analysing the differences in the semiochemical contents of faeces and urine, and trials using selected semiochemicals to overmark urine left by dingoes in pen trials are potential avenues for further investigation. Understanding the chemical composition of dingo urine, and what combinations may constitute the ‘No trespassing’ message, gives only part of the equation. Information on when, why and how dingoes use scent-marking is currently lacking and requires further investigation. A comprehensive insight into dingo use of and response to semiochemicals, together with knowledge of scent-mark chemistry, will maximise the effectiveness of any future use of semiochemicals as a management tool.
... In sum, dogs have no predators in these two environments,, they have a short and rapid reproductive cycle, with an average of five to six pups, the number of dogs increases rapidly as its food supply is assured (Pal 2003). Therefore, the likelihood of the risk of spreading Leishmania will be more increased. ...
Article
Unlabelled: Human visceral leishmaniasis has long been associated with canine leishmaniasis (CL). However, to date, there is no clear information on the status of the disease in dogs in Morocco that could be used by policymakers for the prevention of human cases. This study aims to assess the status of CL in Morocco and its risk factors through an exhaustive literature search. The meta-analysis was performed using RevMan 5.4. The main results showed that the overall prevalence of CL in Morocco is 17% (95% CI: 0.12-0.22), caused by two strains of Leishmania parasite: Leishmania tropica and L. infantum. According to the region, the maximum prevalence was reported in the coastal provinces and in the central part of the country; while, the CL risk was higher in rural area (18% [95% CI: 0.14-0.23]) and at altitude above 1000 m (23% [95% CI: - 0.08-0.53]). Regarding the intrinsic factors, the prevalence of the disease increased with the age of the dog, (30% [95% CI - 0.09-0.68) and the risk was very high in clinically asymptomatic dogs (RR = 2.08 [95% CI: 1.15-3.76]). This study is the first in Morocco indicating the CL prevalence, its geographical distribution and detailing its risk factors. These results are needed to improve management strategies for the canine reservoir of leishmaniasis in Morocco and interrupt the local transmission cycle to humans. Supplementary information: The online version contains supplementary material available at 10.1007/s12639-022-01521-2.
... In sum, dogs have no predators in these two environments, other than being killed by humans, they have a short and rapid reproductive cycle, with an average of ve to six pups, the number of dogs increases rapidly as its food supply is assured (Pal, 2003), Therefore, the likelihood of the risk of spreading Leishmania will be more increased. This reinforces the relevance of designing a dog population management strategy in Morocco based on the promotion of the dog lifestyle in collaboration with government authorities and community and veterinary leaders for the protection of these animals through education and awareness actions, especially to dog owners. ...
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Methods An exhaustive and thorough literature search was conducted to collect data in seven databases and by contacting the authors of unpublished works. The quality of eligible studies was assessed using the Cochrane Collaboration's risk of the bias assessment tool. The meta-analysis was performed using RevMan 5.4. Zotero and R software were used for data management and statistical processing. Results Descriptive analysis shows that 91.9% of dogs in Morocco lived without veterinary care and 75.8% chained. Two strains of Leishmania were found to be involved in both cutaneous and visceral forms of leishmaniasis in dogs: L. tropica and L. infantum, but the latter is the most common. A meta-analysis was conducted, the results showed an overall prevalence of 17% (95% CI: 0.12–0.22). Significant differences were recorded between regions. The maximum values were reported successively in the provinces more bordering the Mediterranean and those in the center of the country. Risk factors were found to be associated with Leishmania in dogs in Morocco. The risk was found to be high in clinically asymptomatic dogs (RR = 2.08 [95% CI: 1.15–3.76; p = 0.02]) with a high prevalence (9% (95% CI [0.06–0.13]; p < 0.00001). Age was strongly associated with the occurrence of the disease, the prevalence is increased with the age of the dog, it is 30% (95% CI [-0.09-0.68]; p = 0.13). The prevalence was also very high in dogs living in rural areas (18% [95% CI: 0.14–0.23]; p < 0.00001) and in altitudes exceeding 1000 m (23% [95% CI: -0.08-0.53]). Conclusion The present systematic review and meta-analysis are the first of its kind in Morocco and indicate that canine leishmaniasis is common in the regions bordering the Mediterranean coast and the center of the country. It will be possible to predict the epidemiological trend of human visceral leishmaniasis. Monitoring and control actions are needed to improve management strategies of the canine reservoir and interrupt the cycle of transmission to humans.
... Similarly, the frequency of urine marking by domestic (McGuire, 2016) and free-ranging (Pal, 2003) dogs was higher in males than in females. Results also show that male dogs are more likely to actively acquire scent information not only about the opposite sex, but also about the same sex than females when encountering an unfamiliar conspecific. ...
Article
Although body sniffing is the most frequent canine interaction in public places, little is known about olfactory behaviors between walking dogs. The aim of this study was to examine the association of dog, human and behavior characteristics with the number of sniffing dogs within a dyad, with the initiation and termination of sniffing, with the first and last area sniffed on the recipient’s body and with the number of areas sniffed on the recipient’s body in walking dogs in public places. We observed 538 dyadic encounters between sexually intact dogs, each led by one owner. We randomly selected one dog from each of 538 dyads, yielding 486 focal dogs actively involved in sniffing for analysis. Interacting dogs were more likely to engage in mutual than single dog sniffing when the owners communicated with each other than when they did not. Dogs more likely initiated sniffing than not when a male dog encountered a female than vice versa. Dogs more likely sniffed the rear than the head as the first area on the recipient’s body when only one dog sniffed the other than when both dogs sniffed each other; and also when they initiated sniffing than when they did not. Dogs more likely sniffed more than one area than only one area on the recipient’s body when they sniffed the head or abdomen than the rear as the first area on the recipient’s body; when both dogs sniffed one another than when only one dog sniffed the other; and also when they initiated sniffing than when they did not. Dogs more likely sniffed the abdomen or rear than the head as the last area on the recipient’s body, when only one dog sniffed the other than when both dogs sniffed one another; and also when they sniffed more than one area on the recipient’s body than when they sniffed only one area. Dogs more likely terminated sniffing than not when an adult dog encountered a puppy than vice versa. In conclusion, dyadic sniffing behaviors between sexually intact dogs, each walking with one owner, were mainly associated with the owners’ communication, the number of sniffing dogs within a dyad, the dog’s sex and age and with the initiation of sniffing.
... However, contrary to expectations, we found no difference in the measured variables between the path-side (0 m), 1 and 8 m lawn samples, indicating that path-side trees and poles act as focal points for dog-deposition, while lawn areas do not. This is likely a function of gender-specific differences in dogs' urinating and scent-marking behaviors (countermarking), with male dogs preferring to urinate directly on trees and poles (overmarking) while females generally do not, instead preferring to urinate near, but not at the same locations as other dogs (adjacent-marking) (Pal, 2003;Lisberg and Snowdon, 2011). ...
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Urban residents and their pets utilize urban greenspaces daily. As urban dog ownership rates increase globally, urban greenspaces are under mounting pressure even as the benefits and services they provide become more important. The urine of dogs is high in nitrogen (N) and may represent a significant portion of the annual urban N load. We examined the spatial distribution and impact of N deposition from dog urine on soils in three urban greenspace typologies in Finland: Parks, Tree Alleys, and Remnant Forests. We analyzed soil from around trees, lampposts and lawn areas near walking paths, and compared these to soils from lawn areas 8 m away from pathways. Soil nitrate, ammonium, total N concentrations, and electrical conductivity were significantly higher and soil pH significantly lower near path-side trees and poles relative to the 8 m lawn plots. Also, stable isotope analysis indicates that the primary source of path-side N are distinct from those of the 8 m lawn plots, supporting our hypothesis that dogs are a significant source of N in urban greenspaces, but that this deposition occurs in a restricted zone associated with walking paths. Additionally, we found that Remnant Forests were the least impacted of the three typologies analyzed. We recommend that landscape planners acknowledge this impact, and design parks to reduce or isolate this source of N from the wider environment.
... Because of their rapid and short reproductive cycle, with an average of five to six puppies, the population number of stray dogs generally grows faster than its food supply. Unlike other wild animals, which defend their territories, when this condition arises, stray dogs disperse freely towards other suburbs [48,49]. The home range of stray dogs is about 4 to 200 km 2 [50], delimiting a dispersion circle with a maximum radius of 10 km. ...
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In Morocco, cutaneous and visceral leishmaniases represent a public health concern. In this opinion paper, we propose to highlight chosen elements that have governed the drastic increase in the incidence of leishmaniases recorded in Morocco during the period between 1990 to 2010 in order to guide the prediction of the expansion of diseases and epidemic events. We highlight that the dispersion of the zoonotic cutaneous leishmaniasis (ZCL) form, caused by the Leishmania major parasite, appears to be closely related to that of its arthropod vector density, which is sensitive to changes in climate. The dissemination of anthroponotic cutaneous leishmaniasis (ACL) was related to an increase in human travel and local tourism during the studied decades. These are linked to economic expansion and infrastructure development. Interestingly, the main ACL foci are spatially aligned with the highways, and their occurrence was synchronized with the building of transportation infrastructure. During the above-mentioned decades, the zoonotic visceral leishmaniasis (ZVL) caused by Leishmania infantum has expanded from its historical northern territories, dispersing outwards in all directions. This spread follows the emergence of hamlets and villages connecting with major cities.
... En ce qui concerne les chiens de compagnie en liberté, on peut supposer qu'ils étaient motivés à naviguer non seulement pour se nourrir, mais aussi pour des raisons de socialité (Pal, 2003(Pal, , 2015Cafazzo et al., 2012 ;Urbanik, 2012). En effet, Cachupín ne pouvait avoir que des interactions très limitées avec les chiens de compagnie en laisse et il a été observé avec des chiens errants. ...
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La ville de Concepción (Chili) présente l'originalité d'avoir en son sein des chiens en liberté. Nous avons analysé les modalités de présence des chiens de cette ville avec une méthodologie qualitative de sciences sociales (prise de notes des observations et interrogation des citoyens). Un premier résultat est qu'un tiers de l'ensemble des chiens de Concepción étaient en liberté. Un deuxième résultat est que l'ensemble des chiens de cette ville relève de six catégories : les chiens en liberté qui comprenaient les chiens errants agressifs ou non envers les humains et les chiens sauvages, ainsi que les chiens de compagnie, les chiens de garde et de mendiants. Nous avons également remis à jour l'inventaire des chiens vivant dans les villes. Ainsi, un troisième résultat est la découverte de trois variantes des catégories existantes : des chiens errants en meute non agressifs envers l'humain et particulièrement proches de lui spatialement et socialement (Sir Perro) ; un chien errant de compagnie vivant dans une rue semblable aux chiens de village, ayant des interactions très étroites avec les gens (Snoopy) ; des chiens de compagnie errants (le vieux basset et Cachupín). Nous concluons que le lien bidirectionnel entre les chiens et les humains explique :-comment ces chiens urbains deviennent partie intégrante de l'identité urbaine de la ville ;-la plasticité comportementale des chiens errants, à travers leur adaptation à des habitats urbains très diversifiés. Nous postulons que la culture humaine ainsi que les diverses zones urbaines à Concepción ont permis cette unique diversité de positionnement sociospatial des chiens (huit catégories) ainsi que leurs adaptations et réinventions socio-spatiales (les chiens traversant des passages pour piétons).
... As dogs urinate to mark territories and communicate [31,32], the impact due to urine can probably not be reduced without harming the social structure of the dog. However, the impact of feces can be significantly reduced if feces would be collected and disposed of properly. ...
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The number of pet animals in the European Union is increasing over the last decades. Few studies with a limited focus in terms of impacts and life cycle stages exist that assess the environmental impacts of dogs. This paper addresses the entire life cycle of a dog. An LCA study on an average dog was conducted considering the pet food and dog excrements, i.e., urine and feces. Fifteen impact categories were analyzed. An average dog has a climate change and freshwater eutrophication potential of around 8200 kg CO2eq and 5.0 kg Peq., respectively. The main contribution to most impact categories over the dog’s life is caused by pet food. Freshwater eutrophication is mainly determined by the dog´s urine and feces. Feces also have a significant contribution to the category of freshwater ecotoxicity. Impacts increase significantly with increasing weight and a longer lifetime of the dog as well as low collection rates of the feces. This LCA study reveals that pet dogs can have a significant environmental impact, e.g., around 7% of the annual climate change impact of an average EU citizen. Optimizing pet food and increasing the feces´ collection rate can reduce the impacts.
... One should take into consideration, of course, that the distribution of sexes and age cohorts were uncontrolled in our sample, which could have an effect on the statistical reliability. On the other hand, there are reports of protective mother dogs (in free-ranging dogs, [57]), with an emphasis on the most frequent agonistic interactions among the adult females [58]. Paul and colleagues [19], however, showed that in free-ranging dogs, grandmothers may provide help with their daughters' puppies. ...
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Socialization with humans is known to be a pivotal factor in the development of appropriate adult dog behavior, but the role and extent of dog–dog interactions in the first two months of life is rarely studied. Although various forms of alloparental behaviors are described in the case of wild-living canids, the social network of companion dogs around home-raised puppies is almost unknown. An international online survey of companion dog breeders was conducted, asking about the interactions of other dogs in the household with the puppies and the pups’ mother. Based on the observations of these breeders, our study showed an intricate network of interactions among adult dogs and puppies below the age of weaning. Alloparental behaviors (including suckling and feeding by regurgitation) were reportedly common. Independent of their sex, other household dogs mostly behaved in an amicable way with the puppies, and in the case of unseparated housing, the puppies reacted with lower fear to the barks of the others. Parousness, sexual status, and age of the adult dogs had an association with how interested the dogs were in interacting with the puppies, and also with how the mother reacted to the other dogs. Our study highlights the possible importance of dog–dog interactions during the early life of puppies in forming stable and low-stress interactions with other dogs later in life.
... It is evident from the foregoing that most of the studies on stray street dogs are related to rabies and other zoonotic diseases (Diehn 2011;Fu et al. 2019;Garcia-Campos et al. 2018;Kato et al. 2003;Rubin and Beck 1982;Villa et al. 2010). The behavior of Indian street dogs has been amply studied (Majumder et al. 2014;Pal 2001Pal , 2003Pal , 2005Pal , 2008Pal et al. 1998aPal et al. , 1998bPal et al. , 1999. However, neither Pal and his colleagues nor Majumder et al. (2014) investigated the circadian behavior of Indian street dogs. ...
Article
This is the first research article that documents circadian variability in behavioral variables, namely resting (Rt) and standing (St) in stray street dogs of Sambalpur city, India. We also estimated the abundance as a function of time of the day and gender in a population of stray dog inhabiting streets of the city. In addition, we determined the association between the behavioral variables and the environmental variables, such as light intensity, sound intensity, temperature and humidity. We determined the abundance of street dogs at 10 hotspots using the photographic capture-recapture technique and Lincoln index equation. In another study, we determined dogs’ density along the three randomly selected routes that connect the beginning (Dhanupali) and end (P.C. Bridge) of the city precincts. We recorded the resting and standing activities of the stray street dogs using still and video cameras at four times of the day continuously over a longitudinal timescale of 72 hours. This study was conducted at four randomly selected dog hotspots. Subjecting the log-transformed time series data to the Cosinor rhythmometry we obtained three different rhythm parameters, such as mesor (M), amplitude (A) and acrophase (Ø) of the rhythm in resting and standing behavior of stray street dogs. We found out both spatial and temporal variability in the behavior of street dogs. The sightings of dogs were always more during the evening and nighttime irrespective of the investigated routes and hotspots. Further, we also observed that the abundance of male dogs was always significantly more as compared with the bitches. A lack of association between two attributes the time of the day and gender apropos the number of sightings of the street dogs was validated by the Fisher’s exact test. Using Pearson’s correlation analysis technique we found a negative relationship between light intensity and resting activity. In addition, we also found a negative association between standing activity and ambient environmental temperature. These findings were complimentary to the observed circadian variability in the resting and standing behavior of the stray street dogs. In conclusion, despite a few limitations, this study documents a statistically significant circadian rhythm in activities of stray street dogs. It also highlights spatial variability in the abundance of dogs on the streets and hotspot localities of the urban Sambalpur. We do have a hunch. It is likely that similar phenomenamight be of common occurrence in many urban areas of the world. These data might also help in addressing street dog menace – one of the major problems the people and administrative authorities of most of the Indian cities and elsewhere worldwide are experiencing since quite long.
... Territory ownership is typically communicated via acoustic (Gardner & Graves, 2005), visual (Van Dyk & Evans, 2007) and/or chemical cues (Pal, 2003), such as scent-marking (Becker, Castelli, Yohn, Spencer, & Marler, 2018;Carazo, Font, & Desfilis, 2008). Numerous studies have focused on the strength or mode of behavioural responses to scent marks (Ferkin, 2015), the type of scent-marking (Petrulis & Johnston, 1997) or which individual (age, sex and social status) responds (Mares, Young, Levesque, Harrison, & Clutton-Brock, 2011). ...
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1.Territoriality is an important process shaping population dynamics, and the defense of a territory is crucial for individuals to increase the duration of territory occupancy and consequently, reproductive success. However, little is known about how the frequency of territory intrusions and subsequent territorial behaviors and aggression by territory owners are affected by external factors, such as population density. This is important, because it can affect mate change (the replacement of one pair member) and dispersal, a key ecological process. 2.The aim of this study was to investigate the behavioral and spatial response of territory owners to intruder pressure as a function of population density in a territorial, monogamous mammal, the Eurasian beaver (Castor fiber). 3.Using a combination of GPS technology, scent experiments, camera trap data, and tail scar observations from an individual‐based long‐term study, we investigated the factors influencing spatial movement patterns by territory owners in response to a simulated intruder, and the factors affecting territory intrusions. 4.We found consistent inverse density‐dependent patterns in territorial behaviors and evidence of conspecific aggression. At lower densities, territory owners detected more simulated intrusions, showed more territorial reactions, and experienced increased conspecific aggression as indicated by tail scars, suggesting increased intruder pressure. 5.Inverse density‐dependent territorial behavior and aggression suggest a potential mechanistic link between inverse density‐dependent natal dispersal and mate change. At low population densities, increased dispersal amplifies intruder pressure, leading to the observed increases in territorial behaviors, conspecific aggression, and previously observed mate turnover, which in turn might increase natal dispersal. Our study demonstrates how population density can affect the behavior and space use of individuals, which is important for territory occupancy and fitness. This article is protected by copyright. All rights reserved.
... 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). ...
Article
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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.
... It is evident from the foregoing that most of the studies on stray street dogs are related to rabies and other zoonotic diseases (Diehn 2011;Fu et al. 2019;Garcia-Campos et al. 2018;Kato et al. 2003;Rubin and Beck 1982;Villa et al. 2010). The behavior of Indian street dogs has been amply studied (Majumder et al. 2014;Pal 2001Pal , 2003Pal , 2005Pal , 2008Pal et al. 1998aPal et al. , 1998bPal et al. , 1999. However, neither Pal and his colleagues nor Majumder et al. (2014) investigated the circadian behavior of Indian street dogs. ...
Article
As of October 2018, about 1.5 billion people around the world use WhatsApp (WA) for messaging, chatting, and for sharing pictures and videos. Indians send “Good Morning” messages religiously to their online friends and acquaintances almost every day. However, we hardly know about the psycho-physiological basis of the behavior characterized by habitual conveyance of “Good Morning,” and “Good Night,” messages using online Apps. In this study, we attempted to analyze at the individual level if this behavior on WA reflects the user’s circadian timing system. We retrieved chats with time stamps from eight subjects and computed periods of the daily “good morning” messaging behavior of each subject. We computed deviations of average periods in each subject from the theoretical circadian period. Single sample two-tailed t-test revealed that none of the average periods of daily message sending habits of eight subjects was statistically significantly different from the theoretical circadian period of the population. All eight subjects, therefore, revealed entrained circadian rhythm in their messaging behavior. This is perhaps the first study to propose that the timings of the social media messages could be used to gauge the status of the endogenous circadian clock of the users of Social Networking Sites (SNSs).
... In line with previous studies [35,36], our results showed that frequency of urine marking is higher in males than in females. Wirant et al. [37] reported that behaviour of spayed and non-oestrous intact females was similar. ...
Article
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.
... I excluded from my studies intact females in proestrus and estrus, periods during the ovarian cycle when specific gonadal hormones increase (Beach et al., 1982). Free-ranging female dogs in estrus exhibited increased rates of urine-marking (Pal, 2003) and privately owned Jack Russell terriers directed more of their urinations at targets in the environment when in proestrus and estrus than when in anestrus (Wirant et al., 2007). Increased urination also characterized beagles spayed as adults and administered exogenous gonadal hormones (Beach, 1974). ...
... The mtDNA was extracted from a swab of the each scat's exterior using QIAGEN (Venlo, Netherlands) DNeasy TM DNA extraction kit following the protocol described in Vynne et al. 59 . Extractions were carried out in a room separate from the lab in which PCR amplifications were performed using dedicated equipment to prevent contamination. ...
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Detection dogs, specially trained domestic dogs (Canis familiaris), have become a valuable, noninvasive, conservation tool because they remove the dependence of attracting species to a particular location. Further, detection dogs locate samples independent of appearance, composition, or visibility allowing researchers to collect large sets of unbiased samples that can be used in complex ecological queries. One question not fully addressed is why samples from nontarget species are inadvertently collected during detection dog surveys. While a common explanation has been incomplete handler or dog training, our study aimed to explore alternative explanations. Our trials demonstrate that a scat's genetic profile can be altered by interactions of nontarget species with target scat via urine-marking, coprophagy, and moving scats with their mouths, all pathways to contamination by nontarget species' DNA. Because detection dogs are trained to locate odor independent of masking, the collection of samples with a mixed olfactory profile (target and nontarget) is possible. These scats will likely have characteristics of target species' scats and are therefore only discovered faulty once genetic results indicate a nontarget species. While the collection of nontarget scats will not impact research conclusions so long as samples are DNA tested, we suggest ways to minimize their collection and associated costs.
... The notion that gonadal hormones affect urinary behaviour also under physiological conditions is supported by the higher proportions of directed urinations during proestrus and oestrus compared with anoestrus (Wirant, Halvorsen, & McGuire, 2007). A noticeable increase in urine marking is also reported for free-ranging female dogs in oestrus (Pal, 2003). In wolves, the influence of gonadal hormones on urinary behaviour is evident, as urine marking is only apparent after puberty, and a seasonal pattern of urine marking rate associated with serum testosterone levels is shown in dominant females (Asa, Mech, Seal, & Plotka, 1990). ...
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Benefits and risks of spaying on the behaviour of female dogs are controversially discussed. Increased aggressiveness and male urinary behaviour were postulated to be the consequence of masculinization after spaying in some female dogs. To investigate if spaying or its timing relative to the onset of puberty may have a masculinization effect, urinary behaviour, that is, frequency of urination, urinary posture and ground scratching after urination were recorded in 58 female Labrador Retrievers during 15 min of a daily walk with their owners. General behaviour of the dogs during the walks was assessed using an owner questionnaire. Data were analysed for age, reproductive status, lifetime of ovary exposure and/or time interval since spaying. Urinary behaviour of intact females (n = 12) and dogs spayed before (n = 17) or after (n = 29) puberty was similar and not influenced by age, lifetime of ovary exposure and/or time interval since spaying. Owners of spayed dogs described more frequent or more intense fear reaction in their animals in response to loud noises, unfamiliar objects approaching on or near the sidewalk, or if they were approached by unknown dogs barking, growling or jumping. In conclusion, we found no evidence of a masculinization effect after spaying on urinary behaviour in female Labrador Retrievers. In contrast to popular belief, gonadectomy did not inevitably result in a behaviourally more stable dog. Extrapolation of our findings from female Labrador Retrievers to other breeds should be performed with caution, as the effect of spaying on behaviour may differ among dog breeds.
... doi: bioRxiv preprint first posted online Jul. 3, 2018; (White, 2008). Since, the dogs are territorial (Pal, 2003) and our surveys were conducted over 132 two or three consecutive days in each village, we assumed the individual populations were 133 closed during survey. Also, since we used non-invasive means to identify dogs, we assumed 134 there was no loss of marks and no change in behavior following the first "capture". ...
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Many mammalian carnivore species have been reduced to small, isolated populations by habitat destruction, fragmentation, poaching, and human conflict. Their limited genetic variability and increased exposure to domestic animals such as dogs place them at risk of further losses from infectious diseases. In India, domestic and feral dogs are associated with villages in and around protected areas, and may serve as reservoirs of pathogens to the carnivores within. Kanha Tiger Reserve (KTR), India, is home to a number of threatened and endangered mammalian carnivores including tiger (Panthera tigris), leopard (Panthera pardus), wolf (Canis lupus), and dhole (Cuon alpinus). It also has more than 150 villages with associated dog populations. We found that dog populations ranged from 14 to 45/village (3.7 to 23.7/km2), and did not vary with village area, human population size, or distance from the core area of KTR, though they all increased between summer 2014 and winter 2015, primarily through reproduction. No dog tested positive for rabies but seroprevalence levels to three other generalist viral pathogens were high in summer (N=67) and decreased somewhat by winter (N=168): canine parvovirus (83.6% to 68.4%), canine distemper virus (50.7% to 30.4%) and canine adenovirus (41.8% to 30.9%). The declines in seroprevalence were primarily due to new recruitments by birth and these were not yet exposed to the viruses. Wild carnivores frequently entered the villages, as shown by tracks, scats, kills and other indicators, and the dogs are known to leave the villages so that encounters between dogs and wild carnivores may be common. We conclude that there is a large population of unvaccinated dogs in and around Kanha Tiger Reserve, with high levels of seroprevalence to pathogens with broad host ranges and these dogs which interact with wild carnivores, therefore posing a high risk of disease spillover to the wild carnivores.
... H1-Infrastructure, the presence of owned and unowned free-ranging dogs is usually associated to human settlements(Acosta-Jamett et al. 2010;Silva-Rodriguez et al. 2010;Gompper 2014;Morters et al. 2014;Villatoro et al. 2016), habituated to feed upon human discarded food and garbage(Pal 2003;Campos et al. 2007;Vanak 2008;Vanak and Gompper 2009b;Atickem et al. 2010;Dias et al. 2013) and using trails and roads to access wild habitats(May and Norton 1996;Fiorello et al. 2006; Silva-Rodríguez et al. 2009; Silva-Rodriguez et al. 2010; Doherty et al. 2015; Moreira-Arce et al. 2015; Sepúlveda et al. 2015; Parsons et al. 2016); H2-Environment, freeranging dogs prefer open habitats such as savannas, pastures and shrublands, since they offer little if any resistance to movement, compared to forest habitat, which can act as a clear barrier to the movement of free-ranging dogs(Meek 1999;Vanak 2008;Lacerda et al. 2009;Vanak and Gompper 2010;Sepúlveda et al. 2015). Open habitats like coastal grounds also harbor several species of waterbirds(Schüttler et al. 2009;Couve et al. 2016), while during fieldwork, livestock could be frequently observed in other types of open habitats like pastures, birds and cattle might be possible prey for free-ranging dogs. ...
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Domestic dogs (Canis familiaris) have turned out to be one of the most common carnivoran species in natural ecosystems worldwide, becoming a major concern for wildlife conservation, particularly on islands. Here, we assessed the effect of infrastructure and the environment on the occupancy probability of free-ranging dogs on Navarino Island, Cape Horn Biosphere Reserve, southern Chile. We collected camera-trap data from 200 sites in a grid around the only major settlement of Navarino Island. Single-season, single species occupancy modeling was used to assess the impact of five infrastructure variables and two environmental variables on the occupancy of free-ranging dog and of six variables on the probability of detection. A total of 4,000 camera-trap days yielded 67 independent photo sequences of free-ranging dogs. Our results provided support for the hypothesis that environmental variables had the most influence on occupancy, when compared to infrastructure variables, while Julian date, survey and animal trail density were the most important predictor variables for detection probability. Free-ranging dogs preferred open habitats instead of forests and habitats at lower elevations. The photographic records further showed interaction between owned/unowned and feral free-ranging dogs as well as reproduction in feral dogs. Dogs were slightly more active at day than at night. Results of the present study demonstrated that there is an urgent need to implement management measurements in order to reduce the numbers of free-ranging dogs in the Cape Horn Biosphere Reserve.
... Consistent with these findings, Lisberg and Snowdon (2011) found that, per dog, frequencies of urination and countermarking in a dog park entryway were higher in males. Previous work on free-ranging dogs and companion dogs in nondog park contexts have also reported greater urinary/marking behaviours in males compared to females (Cafazzo et al., 2012;Pal, 2003;Ranson and Beach, 1985) and in older individuals (Cafazzo et al., 2012;Ranson and Beach, 1985;Wirant et al., 2007;Wirant and McGuire, 2004). ...
Article
This study examines the activity budgets and social behaviours initiated and received by 69 focal dogs in an off-leash dog park for 400 seconds after entry, a time of high activity about which little is known. Using motivationally-neutral labels for social behaviour categories, we describe the frequency of behaviours, and correlations among them. We then examine these relationships in the context of proposed functions for some behaviours in dogs, in terms of information gathering and communication, including visual and tactile signalling. Time spent with other dogs decreased rapidly over the visit, and much of this early interaction involved greeting the park newcomer. Snout-muzzle contact behaviours were ubiquitous, while other behaviours were rarely observed, including aggressive behaviours. Correlations among certain non-contact behaviours initiated and received by focal dogs are consistent with their function as visual signals that may influence the continuation and form of social interactions, and their possible role in social mimicry (i.e., play bow and pull-rear away). Age, sex, and number of dogs present in the park influenced specific aspects of dogs' activity budgets, and a few behaviours. This ethological study provides fundamental data on dog social behaviour in dog parks, about which surprisingly little has been published.
... This finding is consistent with our previous suggestion that some mature males may initially respond to the challenging conditions of shelter life by temporarily reverting, in whole or in part, to the juvenile leanforward posture in which no hindlimb is raised (Gough and McGuire, 2015). Percentages of urinations with a raised hindlimb were somewhat lower in adult males at shelters (73% at the Tompkins shelter and 86% at the Cortland shelter; Gough and McGuire, 2015) than adult males observed under other conditions, such as laboratory colonies or free ranging (94%-99%; Sprague and Anisko, 1973;Beach, 1974;Bekoff, 1979;Pal, 2003;Cafazzo et al., 2012). Also, temporary return to the lean-forward posture has been reported for adult male dogs in fearful situations (Berg, 1944;Martins and Valle, 1948). ...
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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.
... This result was agreed with those recorded by (Betty, 2016;Sharon and Betty, 2004), which found that dog, urinate more frequently than bitches and had high tendency to direct urine at specific objects in the environment a scent marking. Also (Pal, 2003) found that free-ranging dogs with a high tendency to direct urine for near territory boundaries while bitches marked at nest sites, so location with strong influence on male and female urinary behavior of free-ranging dogs. Also (Gese and Ruff 1997) reported that the most common posture in female squat rising, which found that lifting the leg during urination in unusual among bitches and during estrus. ...
... Conversely, Although ubiquitous, here we concentrate on free-ranging Domestic Dogs in India and Australia, which have abundant and widespread populations of them Pal, 1999;West, 2008). The Indian pariah dog is sometimes referred to as A C C E P T E D M A N U S C R I P T (Bonanni et al., 2010;Fox et al., 1975;Pal, 2003Pal, : 2008. ...
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The roles of the 37 species in the family Canidae (the dog family), are of great current interest. The Gray Wolf is the largest canid and their roles in food webs are much researched, as are those of Domestic Dogs, Coyotes and Red Foxes. Much less is known about the other canid species and their ecological roles.Here we describe general food web theory and the potential application of network theory to it; summarise the possible roles of predators in food webs; document the occurrence, diet and presumed functions that canids play in food webs throughout the world; give case studies of four threatened canid species of top, middle and basal trophic positions and six anthropogenically affected species; and identify knowledge limitations and propose research frameworks necessary to establish the roles of canids in food webs.Canids can be top-down drivers of systems or responsive to the availability of resources including suitable prey. They can be affected anthropogenically by habitat change, lethal control and changes to basic resource availability. They can be sustainable yield harvesters of their indigenous prey or passengers in complex ecosystems, and some are prey of larger canids and of other predators. Nevertheless, the roles of most canids are generally poorly studied and described, and some, e.g. Gray Wolves, Coyotes and Australian dingoes, are controversial. We advocate mensurative and experimental research into communities and ecosystems containing canids for a quantitative understanding of their roles in food webs and consequent development of better management strategies for ecosystems.
... Agonistic interactions can be intense (Drews 1993), especially during territorial contests, which can lead to Communicated by A. Pilastro Electronic supplementary material The online version of this article (doi:10.1007/s00265-016-2260-6) contains supplementary material, which is available to authorized users. (Gese and Ruff 1997) and aggressive propensity (Pal 2003). Also in fishes, transmission of chemical information via urine plays a role in agonistic interactions (Hirschhauser et al. 2008;Maruska and Fernald 2012). ...
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The communication of aggressive propensity is an important component of agonistic interactions. For this purpose, animals use different sensory modalities involving visual, acoustical and chemical cues. While visual and acoustic communication used in aggressive encounters has been studied extensively in a wide range of taxa, the role of chemical communication received less attention. Here, we studied the role of chemical cues used during agonistic interactions of territory owners in the cooperative cichlid Neolamprologus pulcher. During staged encounters, we allowed either visual and chemical contact between two contestants or visual contact only. As chemical information in this species is most likely transferred via urine, we measured urination patterns using dye injections. Furthermore, we recorded aggressive and submissive behaviours of both contestants in response to the experimental treatment. Fish that had only visual contact with each other significantly increased their urination frequency and showed more aggressive displays compared to fish with both visual and chemical contact. Furthermore, appropriate agonistic responses appear to be dependent on available chemical information. This indicates that N. pulcher actively emits chemical signals to communicate their aggressive propensity via urine. Chemical communication thus plays a crucial role in multimodal communication of aggression in these fish, which highlights the need of studying the role of chemical communication during agonistic encounters in general, even if other signals are more obvious to the human observer.
... We obtained pooled dingo urine collected from a large pack of dingoes at the Australian Dingo Conservation Association in April 2013. Canines often urinate to mark territories (Pal 2003). A pooled sample is more likely to contain a greater range of semio-chemicals than samples collected from individual dingoes. ...
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Southern hairy-nosed wombats (Lasiorhinus latifrons) are fossorial marsupials that live in large burrow systems where their digging behaviour brings them into conflict with agriculture. In the absence of any available control options, non-selective culling is the primary mode of wombat management. This approach is contentious and has unknown implications for long-term wombat conservation. Predator scents, however, have been effective in altering behaviours of some herbivores and may offer a non-lethal alternative to culling if they discourage wombats from burrowing in perceived problem areas. Therefore, we trialled two dingo scents (faeces, urine) over 75 days to determine whether these scents would deter wombats from repopulating collapsed burrows. Ten inhabited single-entrance burrows were excavated over three days (to allow time for inhabitants to exit), collapsed and then filled in. Five burrows, separated by at least 200 m, were used for dingo scent treatments (three urine; two faeces) and three burrows, separated by the same distance, served as negative controls (unscented), along with two ‘farmer-monitored’ active controls (dog urine and a dingo carcass). We used a rank-sum score to assess wombat activity: scratching was scored with a value of (1), digging (2), and recolonisation (5), with each value reflecting total energy and time spent in the vicinity of the treatment. We fitted Generalised Estimating Equations (repeated-measures, Fisher Method) to explain variation within, and across, treatment and control burrows. Within 20 days, all 10 sites had signs of wombat activity that ranged from fresh digging, to fully functional burrows. Among the five treatment sites, scratching and tracks identified wombats as being present, but they did not dig. After 75 days, the five sites treated with dingo scents had minimal activity and no new burrows, while wombats recolonised all control burrows. Though we used only 10 burrows for this preliminary study, our findings suggest the need for further testing of dingo scents as a tool for dissuading wombats from digging and recolonisation of collapsed burrows. This represents a novel use for a predator scent, in that prey may remain in the vicinity near the deterrent, but curb problematic behaviours of economic consequence.
... In feral dogs, reproduction is not linked with dominance. Pal et al. (1999Pal et al. ( , 2003Pal et al. ( , 2005 could neither find clear displays of dominance or submission in their group of feral dogs nor frequent aggressive encounters between females. They found little reproductive suppression in females, and they found that infanticide only occurred rarely. ...
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A dominance hierarchy is an important feature of the social organisation of group living animals. Although formal and/or agonistic dominance has been found in captive wolves and free-ranging dogs, applicability of the dominance concept in domestic dogs is highly debated, and quantitative data are scarce. Therefore, we investigated 7 body postures and 24 behaviours in a group of domestic dogs for their suitability as formal status indicators. The results showed that high posture, displayed in most dyadic relationships, and muzzle bite, displayed exclusively by the highest ranking dogs, qualified best as formal dominance indicators. The best formal submission indicator was body tail wag, covering most relationships, and two low postures, covering two-thirds of the relationships. In addition, both mouth lick, as included in Schenkel's active submission, and pass under head qualified as formal submission indicators but were shown almost exclusively towards the highest ranking dogs. Furthermore, a status assessment based on changes in posture displays, i.e., lowering of posture (LoP) into half-low, low, low-on-back or on-back, was the best status indicator for most relationships as it showed good coverage (91% of the dyads), a nearly linear hierarchy (h' = 0.94, p
Chapter
Dogs, together with humans, are one of the most successful species on the planet. In the Western world, dogs usually live as companions and often take on important roles for their human partners. However, a larger proportion of dogs worldwide are free-ranging – living in the human environment, but otherwise making their own decisions. In this chapter we summarize the various aspects of the socio-ecology of these dogs. The discordant reports, going from solitary to structured packs, likely reflect the flexibility and adaptability of this species as well as possible genetic differences between populations of FRDs. The high variation in sociality begs the question of which socio-ecological factors, such as food distribution and abundance, interdependence, closeness to humans, etc., may be affecting the dogs’ social structure. Where packs do form, the internal structure remains still scarcely understood. It appears that, in most cases, groups are composed of multiple males and females, but mostly due to the absence of genetic data and/or long-term studies, it remains unclear whether the pack members are related with each other or not. Both males and females are largely promiscuous, and parental care seems to be mostly provided by the mother, with varying support from putative fathers and other (related?) females. Dog packs collectively defend established territories, marking and engaging in defensive aggression against other packs.
Chapter
Behavior can change as a result of medical problems or physiological changes, and behavior changes are likely to be the first signs of stress, disease, and poor welfare in any animal. If shelter operations, behavior, and/or medical staff identify behaviors that may have an underlying medical cause, they can be addressed immediately, relieving suffering and increasing the adoptability of the animal. Conversely, if medical conditions that cause or exacerbate problematic behaviors are missed, time may be wasted on training or attempted behavior modification, thus prolonging suffering and time spent in the shelter. Only by safeguarding both physical and emotional health can we improve overall quality of life for animals in our care, facilitate their placement in homes, and help prevent their return to the shelter.
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The maned wolf ( Chrysocyon brachyurus ) is an induced ovulator. Though the mechanism of ovulation induction remains unknown, it is suspected to be urinary chemical signals excreted by males. This study assessed volatile organic compounds (VOCs) in weekly urine samples across 5 months from 13 maned wolves (6 intact males, 1 neutered male, 6 females) with the goal of identifying VOCs that are differentially expressed across sex, reproductive status, and pairing status. Solid-phase microextraction (SPME) and gas chromatography-mass spectrometry (GC-MS) were used to extract and separate VOCs that were identified via spectral matching with authentic standards, with spectral libraries, or with new software that further matches molecular fragment structures with mass spectral peaks. Two VOCs were present across all 317 urine samples: 2,5-dimethyl pyrazine and 2-methyl-6-(1-propenyl)-pyrazine. Fifteen VOCs differed significantly ( Adj . P < 0.001 and |log 2 fold change| >2.0) between intact males and females. Using partial least squares-discriminant analysis, the compounds with the highest importance to the sex classification were delta-decalactone, delta-dodecalactone, and bis(prenyl) sulfide. Sixty-two VOCs differed between intact males and the neutered male. Important classifier compounds were 3-ethyl 2,5-dimethyl pyrazine, 2-methyl-6-(1-propenyl)-pyrazine, and tetrahydro-2-isopentyl-5-propyl furan. Several VOCs established as important here have been implicated in reproductive communication in other mammals. This study is the most robust examination of differential expression in the maned wolf thus far and provides the most comprehensive analysis of maned wolf urinary VOCs to date, increasing the sample size substantially over previous chemical communication studies in this species. New data analysis software allowed for the identification of compounds in the hormone-producing mevalonate pathway which were previously unreported in maned wolf urine. Several putative semiochemicals were identified as good candidates for behavioral bioassays to determine their role in maned wolf reproduction, and specifically in ovulation induction.
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1. Chemical sensing in vertebrates is crucial in their lives, and efforts are undertaken towards deciphering their chemical language. Volatile organic compounds (VOCs) is a group of chemicals believed to play an essential role in a wide variety of animal interactions. Therefore, understanding what animals sense themselves and untangling the ecological role of their volatile cues can be accomplished by analysing VOC emissions. A Proton-Transfer Reaction Time-of-Flight Mass Spectrometer (PTR-TOF-MS) is an instrument that measures VOCs in real-time in an air sample. Since this technique acts as a hyper-sensitive ‘nose’ it has a similar potential in deciphering the chemical language of vertebrates. 2. Here, we validate the use of PTR-TOF-MS as a tool to measure VOCs from vertebrates, which in turn will help resolve vertebrate interactions through VOCs. The instrument monitors and records the full spectrum of VOCs emitted by an individual with a high accuracy and low detection limit, including transient VOC emissions. We propose and test diverse measuring configurations that allow for measurement of VOC emissions from different vertebrates and their exudates: full body, specific parts of the body, urine and femoral pores. In addition, we test configurations for sudden and short-lasting processes as VOCs emitted during adder skin shedding as well as the emissions of skin secretions upon mechanical and physiological stimulation in amphibia. Our configurations work in tandem with Gas Chromatography Mass Spectrometry (GC-MS) to allow compound structure verification. 3. We discuss the configurations and methodologies used and conclude with recommendations for further studies, such as the choice of chamber size and flow. We also report the results of the measurements on vertebrates —that are novel to science— and discuss their ecological meaning. 4. We argue that PTR-TOF-MS has a high potential to resolve important unanswered questions in vertebrate chemical ecology with great adaptability to a wide range of experimental setups. If combined with a structure verification tool, such as GC-MS, the creative deployment of PTR-TOF-MS in various future study designs will lead to the identification of ecologically relevant VOCs.
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Large carnivores are important for biodiversity conservation, constituting an essential component in maintaining integrity and stability of ecosystems. In consequence, carnivores’ conservation and the development of suitable management plans for them are crucial for the maintenance of whole ecosystems. These plans, to be successful, need complete information about species, which includes the study of their behaviour. However, the study of large carnivores is widely known for being very complicated, mainly because of their evasive behaviour and large home ranges, remaining most of them hardly studied, and many aspects of their ecology and behaviour being still insufficiently unexplored. This study focuses on jaguars (Panthera onca) and pumas (Puma concolor), the largest felids of the Neotropics. Both are considered umbrella species, as they cover all the ecological requirements of a community; and key species because of their direct and indirect influences over the different animals and vegetation species, maintaining the stability of ecological communities. They are also considered as flagship species, as they are easy attractors of environmental campaigns. Neither of this species is classified as threatened globally by the IUCN; however, they are not exempt from human pressures derived from population growth, for example, which triggers habitat fragmentation and loss or hunting pressure, with devastating effects on biodiversity. The general decrease of jaguar and puma population requires a search for alternatives to conflicts where they are involved. The understanding of the ecological needs of both species and their interaction with the ecosystem will result in the improvement of their own conservation status, but also of the rest of species within the community. A good option for the study of these elusive species is non-invasive methods, which usually allow the survey of most part of the population, being a suitable method for the investigation of carnivores ́ ecology, behaviour and population status. Specifically scats are easy surveyed and they present a lot of ecological and territorial uses for individuals, making their collection and analysis a powerful tool for the study of large carnivores ́ ecology. Furthermore, the importance of scats-based studies has greatly increased with the advances in genetic techniques, allowing researchers to identify species, sex and individuals from DNA extracted from them. This thesis is specifically centred in the study of the marking behaviour of jaguars and pumas in different areas of the Neotropics. With the study of the scat-marking patterns we aim to improve the knowledge about the behavioural ecology of these large carnivores. First, and to be able to interpret correctly the results of the scat spatial analysis, it is necessary to know the spatial behaviour of the species. Until these days, few studies have analysed the spatial behaviour of pumas and jaguars. Besides, existing studies are heterogeneously distributed along the geographical range of the species, with large areas remaining unstudied. Some aspects of the spatial ecology of the species are also unknown (e.g. there is hardly any information about the daily movements and dispersal behaviours of jaguars) and the methodology is highly different among studies, both in the field and in the posterior analyses. Consequently, and to deal correctly with the scats' spatial distribution analysis, in the first chapter we reviewed all the available studies about the spatial ecology of jaguars, pumas and ocelots to try to clarify and summarize all the information available about the different aspects of their spatial ecology. Here, we decided to also include the ocelot, the third largest felid of the Neotropics, as we considered important to have as much available information as we could about the largest felids of the Neotropics. This review showed that pumas presented on average larger home ranges than jaguars and ocelots. Three variables, sex, ecoregion and latitude influenced the home range size of all three species; whereas only sex and latitude showed significant influence in the daily net displacement of ocelots. Results regarding territoriality showed that intersexual range overlap was higher than intrasexual in jaguars and pumas. Previously to analyzing the scats spatial distribution, it is necessary to take into account different variables which could affect the scats surveying and produce misleading results. In chapter 2, we analysed how different climatic and vegetation variables affect the scat detection rate in different areas of the Neotropics. Although biases and interpretational errors have been detected for laboratory and data analyses when using scats for ecological studies, biases related to scats sampling have rarely been discussed. The time that a scat remains recognizable in the field is almost unknown, and it can be an important source of error in many studies. In fact, our study showed that total precipitation and mean temperature of the three months preceding the survey negatively influenced scat detection rate; nevertheless, neither detection method, nor vegetation density and height, nor ecoregion did show any significant effect. Studies based on the scat detection rate of jaguars and pumas may be biased by the influence of some of the mentioned factors affecting scat persistence. Investigators should minimize the biases induced by these variables taking into account, specially, precipitation levels and temperature of the areas surveyed. Once we studied the variables influencing scat detection rate in our surveyed areas, in chapter 3 we proceeded to explore the scats' spatial distribution pattern. It is known that scats are usually deposited over prominent and conspicuous sites or at strategic points on trails near the central area; especially at nearby junctions, points which are frequently remarked. These signals are used for leaving long term messages, such as territory marking and maintenance, individual recognition or signaling feeding places or sites with valuable resources. Here, we examined several aspects of jaguars and pumas spatio-temporal marking behaviour based on scats. Specifically, we examined the frequency of remarking at small and medium spatial and temporal scales, analyzing if scats were randomly deposited along their territory and evaluating if deposition points were mainteined over time. We observed that both species showed a remarking behaviour at specific places, confirming the use of scats as markign signals; however, jaguars remarked with a higher frequency and deposited the scats in a more aggregated way than pumas. This would suggest the dominant role of jaguars versus the subordinate species, pumas, which would try to avoid encounters with jaguars. Finally, in chapter 4, an to complement the knowledge available about the marking behaviour, we used scrapes data, as this is a known signal of communication among individuals. A couple of studies exist about the scrape-marking behaviour of jaguars and pumas, although these studies are mainly focused on generalities and they do not delve into specific aspects of this behavioural trait. This is partly understandable given that the interpretation of scrapping behaviour present some difficulties, one of them being the identification of the particular species that made the scraps. Only in the cases that there are fresh scats or urine over it, the owner of the mark can be identified and thus, the specific scrape can be assigned to a single species. Our results showed that most of the scrapes were found in car tracks, and they were usually located in the edge of the track; specifically, felids make scrapes on sites mainly covered by leaves and located along narrow paths, clean and rarely used. When comparing between areas, scrapes were located mainly in the centre of the path in areas only with pumas, in the centre and in the edge in areas with a similar number of jaguars and pumas, and in the edge in area mainly dominated by jaguars. This could indicate the existence of different scrape marking behaviours in the two studied species, although further studies are needed to confirm these findings. In conclusion, there is still much work to be done in order to obtain a detailed knowledge about jaguars and pumas marking behaviour, and about the use of the space to deposit scats and scrapes for intra and interspecific communication. However, the presented analyses of the marking behaviour (scats and scrapes) of the largest felids of the Neotropics is a first step towards the full comprehension of the spatial ecology of these two species facing increasing human pressures, and highlight the importance of their interacting patterns.
Chapter
This chapter reviews olfactory behavior in the Carnivora, concentrating on the family Canidae. It is divided into three main sections. First, aspects of the olfactory system that may contribute to the canids' acute sense of smell are considered. Second, the olfactory behavior of canids is discussed, and, finally, the applications of dogs' olfactory abilities are reviewed. The chapter uses selective examples to illustrate olfactory behavior in canids, and highlights issues which require further study in order to fully understand the role of odors for these species. The olfactory system of the canids is basically similar to that found in all vertebrates. It has been speculated that individually identifiable odors may originate from the faeces, urine, feet, anal sacs/gland, supracaudal gland, and skin. The olfactory abilities of dogs, combined with their trainability, have led to their use to detect and discriminate odors in forensic, economic, conservation, and health and medical applications.
Article
Dogs are a popular pet in the United Kingdom and walking a dog is widely recognized as an important part of dog ownership. A number of different restraints can be used when walking dogs on leashes such as collars and harnesses. Previous research has examined the behavioral effects of walking dogs on head and neck collars. Harnesses are often anecdotally proposed to be more beneficial to dog welfare than other alternative restraints, however to date the effects of walking dogs on harnesses have not been investigated. The aim of this study was to determine the behavioral responses of dogs walked on neck collars or harnesses. The broader purpose of this study was to examine if the type of restraint worn causes stress in dogs. In order to explore this, a within-subject counterbalanced design was used. Thirty privately owned dogs were recruited within two groups (each group: n=15); those previously walked on a harness and those previously walked on a neck collar. Dogs were walked for 20 minutes each while behavioral indicators of stress were recorded. Post this trial, owners were given the alternative walking restraint and returned a week later to perform a second 20 minute walk. Behavioral indicators were again recorded. No significant differences were found between behaviors shown by dogs when walked on either collar or harness. However dogs with a history of being walked on a collar showed increased low ear position. This may suggest that these dogs are more stressed however due to the lack of support from the other stress indicators, motivations, such as indicating appeasement toward their owners, should also be considered. These findings suggest that, at least for the specific harness and collar trialed, neither neck collars nor harnesses are eliciting stress in dogs. However, future research determining the long-term effects of neck collar and harness use would be beneficial.
Chapter
The maned wolf (Chrysocyon brachyurus) is one of only two known canids exhibiting induced ovulation, a phenomenon that may be controlled by semiochemicals. This study employed headspace solid-phase microextraction with GC–MS coupled with the recursive use of Agilent’s MassHunter Workstation and Mass Profiler Professional software to investigate the volatile organic compounds (VOCs) from monthly urine samples of 11 maned wolves. The goals were to find compounds that differ between males and females as a first step toward semiochemical discovery and to create a list of compounds found commonly across samples to form the basis of a control mixture for use in behavioral bioassays. Ten compounds were found to differ significantly between males and females (p < 0.001 and fold change >3.0): ψ-diosphenol, 1,3-di-tert-butylbenzene, and 2,4-dimethyl-1-heptene were respectively 3000, 600, and 85 times more abundant in males than females. Butanoic acid was unique to female maned wolves and nonanoic acid was 19 times more abundant in females than males. Twenty-five compounds were identified in >98 % of the samples. Several of these compounds have been previously identified in maned wolf urine and some have been reported as semiochemicals in other mammal species. The analysis demonstrates that HS–SPME–GC–MS combined with automated data processing can successfully shorten the list of compounds that require manual inspection and identification. The use of a recursive software workflow largely automates the search for maned wolf candidate semiochemicals, enabling an intense manual focus on compounds of interest.
Chapter
Part 1: Intrafamilial and Extrafamilial AggressionPart 2: Intraspecific Aggression
Chapter
This chapter reviews evidence for conceptual learning in nonprimate mammals. It shows that there can be no reasonable doubt that highly complex stimuli in the visual, olfactory, and auditory domains can all be reliably discriminated by mammals from a range of groups. The evidence that such discrimination is at the conceptual level (i.e. that discriminably different stimuli are all assigned the same response), rather than involving a simple trigger feature or labelled line, is less conclusive: in some cases it is simply absent; in others it is indirect or inferential. However, in at least some cases, variation in the stimuli has been demonstrated directly, and the data give no reason to doubt that category discrimination is fully possible in these groups.
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This study details the basic ecological behaviors of activity, range and social interaction of 15 individual pets that are permitted varying degrees of freedom to roam without human supervision. The degree of restraint provided by the owner significantly influences the pet's range and interaction with people and other dogs. Pets that are provided with no supervision behave more like un-owned strays than those that are only occasionally permitted to run free.Knowledge that pet dogs roam more extensively the more time they are kept unrestrained may encourage dog owners actively to confine their pets and obey leash laws. That is, control laws will appear less arbitrary and more consistent with the best interests of the community. In addition, the relatively small ranges of pets that are only occasionally permitted freedom may be utilized by animal-control personnel as a management tool; it is more efficient to coax an animal back home, rather than capture it in the hopes it will be retrieved by the owner.
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Different aspects of the etho-ecology of the stray dog population in Valencia (Spain) were investigated. Densities of between 127 and 1304 stray dogs km−2 overlap with density estimates reported for other populations. A male:female ratio of 2:1 is also in agreement with earlier studies. Behavioral observations revealed that these dogs will occasionally form groups with dominance hierarchies and communal defense of a territory. From the stability of these groups, long-term affiliative bonds apparently exist among group members. This finding conflicts with the accepted notion that urban stray dogs are asocial and do not form stable social groups. Methodological problems may invalidate earlier claims that urban stray dogs are asocial animals. This is due to improper use of a Poisson model for assessing social organization. It is suggested that stray dogs possess, like most canids for which an adequate data base exists, remarkable behavioral plasticity allowing them to adjust their social system to prevailing ecological constraints.
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With the aid of x-ray ventriculography, lesions were placed in the medial preoptic-anterior hypothalamic (MP-AH) area of 5 adult male pygmy goats. The lesions resulted in a marked decrement in ejaculatory responses, as they have in previous studies on rats, cats, dogs and rhesus monkeys. The results illustrate the apparent universal effects of MP-AH lesions on copulatory behavior in mammalian species. Male sex-typical non-courtship responses of flehmen, self-enurination and penis licking were not altered by the MP-AH lesions. Since the non-courtship responses are displayed most frequently when male goats are sexually aroused, the results point to a specific effect of the lesions in disturbing a male's copulating performance while sparing other expressions of sexual arousal.
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Discusses variation in Felis catus social behaviour, distinguishing between: 1) populations in which adult females are generally solitary and those where they are gregarious; 2) variation in group size and social structure, and 3) individual variation within and between age and sex classes in terms of the nature of social relationships. Specifically, note is taken of the significance of spacing, encounters, use of social odours, group size and structure, and social dynamics. Attention is paid to the nature and consequences of the social hierarchy that is established in farm cat societies, which varies considerably between populations. -S.J.Yates
Article
Population size and density, age structure, survivorship patterns, sex ratios, and social organization of urban, rural, and feral dog (Canis familiaris) populations were examined in Cd. Juarez, Mexico (urban site) and on the Navajo reservation (rural and wild sites) between June 1983 and December 1984. Urban and rural dogs were less social than expected whereas feral dogs characteristically lived in packs. Seasonal variation in the structure of feral dog packs was influenced by reproduction, both directly (pups born into the pack) and indirectly (pregnant females may temporarily emigrate form the pack to give birth).
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.
Article
This paper reports aspects of a long-term study (1975-84) of the ecology, social organisation and behaviour of dingoes, Canis familiaris dingo, on the lower Fortescue River in Western Australia. In all, 170 dingoes were fitted with radio-collars and tracked from aircraft. Dingoes were sighted during 59% of the 13 618 occasions that they were being radio-tracked during the day. Radio-tracking yielded 31 229 daytime and 3016 night-time locations of radio-collared dingoes. The average duration of radio contact with 146 dingoes was 9 months (range 1-35 months). Dingoes were most active around sunrise and sunset, moderately active during the night, and least active during the heat of the day. Travelling (local meandering and more purposeful movement) was the most commonly witnessed activity. Levels of scent-marking (raised-leg urination and ground-scratching), howling and general activity increased over the 2-3 months prior to the mating period, suggesting that dingoes may have a long pro-oestrus (1-2 months). Whelping took place from mid-May to mid-August (mean date 18 July). The characteristics of natal dens are described. The pattern of activities associated with pup-rearing, including alloparental behaviour, closely followed that of related canids.
Article
The dispersal of free-ranging dogs, Canis familiaris, from the town of Katwa, West Bengal, India, was studied from January 1993 to December 1996. Between January 1993 and September 1996, 315 pups were observed from 64 litters. Pups were born between October and March each year, with a peak between November and January. The mortality rate was 68% during the first 4 months, with 102 individuals surviving to the juvenile stage (4 to 12 months). In case of juveniles, the rate of dispersal was 39.29%, whereas, in case of adults it was 23.33%. Mean (±S.D.) home range size for the non-dispersing dogs was 4.8 (±1.7) ha and for the dispersing dogs was 8.4 (±1.7) ha. Moreover, there were significant seasonal variations in the home range sizes of both non-dispersing and dispersing dogs. Juvenile males were the predominant dispersers. Dispersal occurred in all seasons and dispersal rates did not differ between seasons. However, during late monsoon (September to November), dispersal was greater (P
Article
Inter- and intra-sexual behaviour of free-ranging dogs were studied in the natural environment. Both males and females differed in their degree of attractiveness to the opposite sex. There was a positive correlation between the number of males associated with a particular oestrous bitch and the duration of each association. The males preferred by the oestrous bitches enjoyed the highest rates of copulation, but the non-preferred dominant and powerful males occasionally mated forcibly. The young adults were more likely to copulate successfully than the old adults. There was a negative correlation between the number of males present in an association and the number of successful copulators. Occasionally, copulations were performed by opportunist males after being present only a short time. In spite of selective receptivity, a few bitches mated promiscuously. Sometimes the oestrous bitches showed different types of solicitations to their preferred males. In the courting places, aggressiveness of males to other males was higher than the aggressiveness of the females towards other females. Both male–male and female–female mountings from side or rear were observed in this study.
Article
Carnivores are complex creatures living complex social lives in which order is maintained by the transmission of information between individuals. Sometimes the signals are passed visually, sometimes by sound, and very often by odor.
Article
Carnivores use various scent-marking methods. Semi-feral domestic cats (Felis silvestris catus) were observed to use the same means as their wild counterparts. Adult males performed most urine spray marking. Cats scratched tree bark, producing a visual mark, and probably used trees both as markers and for claw sharpening. Most scratching trees were located along frequently used paths rather than along territorial boundaries or scattered randomly throughout a home range. Bark consistency affected the tree species that were scratched, with soft bark preferred. Although deposition of faeces and urine was recorded, there was no clear evidence for their use as territorial markers; cats primarily eliminated away from the core area of the home range. Most faeces were buried, although exposed deposits were also observed. Cats also rubbed against objects, probably using glandular secretions from the face and tail areas to scent mark. Males rubbed objects more than females, and males scent marked more. Individual males may use different means of scent marking. Scent marking in this study supports the idea that cats do not defend territories, instead patrolling and reinforcing marks throughout a looser home range. The suggestion has been made that different forms of marking may serve separate signalling functions.
Article
Seven major types of sampling for observational studies of social behavior have been found in the literature. These methods differ considerably in their suitability for providing unbiased data of various kinds. Below is a summary of the major recommended uses of each technique: In this paper, I have tried to point out the major strengths and weaknesses of each sampling method. Some methods are intrinsically biased with respect to many variables, others to fewer. In choosing a sampling method the main question is whether the procedure results in a biased sample of the variables under study. A method can produce a biased sample directly, as a result of intrinsic bias with respect to a study variable, or secondarily due to some degree of dependence (correlation) between the study variable and a directly-biased variable. In order to choose a sampling technique, the observer needs to consider carefully the characteristics of behavior and social interactions that are relevant to the study population and the research questions at hand. In most studies one will not have adequate empirical knowledge of the dependencies between relevant variables. Under the circumstances, the observer should avoid intrinsic biases to whatever extent possible, in particular those that direcly affect the variables under study. Finally, it will often be possible to use more than one sampling method in a study. Such samples can be taken successively or, under favorable conditions, even concurrently. For example, we have found it possible to take Instantaneous Samples of the identities and distances of nearest neighbors of a focal individual at five or ten minute intervals during Focal-Animal (behavior) Samples on that individual. Often during Focal-Animal Sampling one can also record All Occurrences of Some Behaviors, for the whole social group, for categories of conspicuous behavior, such as predation, intergroup contact, drinking, and so on. The extent to which concurrent multiple sampling is feasible will depend very much on the behavior categories and rate of occurrence, the observational conditions, etc. Where feasible, such multiple sampling can greatly aid in the efficient use of research time.
Article
Urination and defaecation patterns of free-ranging coyotes (Canis latrans) were studied in the Grand Teton National Park, Jackson, Wyoming, for two years. The vast majority of urinations by adult males and females were involved in ‘marking’, and differentiating between ‘marking’ and ‘elimination’ may not be necessary. Our results may be summarized as follows: (1) raised-leg urinations (RLU) performed by males were most frequently used in marking. (2) Females marked throughout the year using the squat (SQU) posture. (3) Snow tracking and reading snow sign resulted in a gross underestimate of the relative frequency of SQU's and a large overestimate in the relative frequency of defaecations (DEF) when compared to results obtained by direct observation. (4) There was sexual dimorphism for the contexts in which marking occurred. Overall, marking by males was associated with courtship and mating, with travelling, and with aggression. Marking by females was associated with the acquisition and possession of food and with the denning season. (5) Marking rates per coyote increased in groups larger than two animals. (6) RLU marking rates were greatest in areas of high intrusion when compared to denning areas and areas in which non-group members infrequently tresscent odours are important in orienting individuals in space but do not represent in and of themselves barriers to movement.
Article
Over a period of 7 months, a residential area of Berkeley, California, was surveyed at hourly intervals throughout the day and night for free-ranging dogs, dogs with people, and cats. The animals' locations and behaviours were recorded. The highest frequency of sightings of free-ranging dogs occurred at 07.00 h, with a secondary peak at 17.00 h. The abundance of dogs with people followed a similar pattern except that the morning peak occurred 2 h later. Cats were most abundant at night between 18.00 and 04.00 h. The observable number of free-ranging dogs increased with increasing temperature to a maximum at 23° C and then declined with higher temperatures. The number of cat sightings was fairly constant with increasing temperature up to 17° C then declined with further temperature increase. Free-ranging dogs were sighted most frequently on private property. Street and side-walk use was greatest in the mornings; free-ranging dogs tended to travel in the morning and rest as the temperature increased later in the day. Most dogs were sighted within 1 or 2 blocks of their homes. The home ranges of 8 dogs are illustrated and described. These showed considerable individual variation, but generally had dense core areas and sparse travel areas (mean corridor home range = 1.74 hectares). Most free-ranging dogs were solitary and social groupings occurred only randomly. Despite their apparent lack of sociability, the dogs did not exhibit any signs of territoriality and agonistic encounters were never observed.
Article
Free-ranging domestic dogs (Canis familiaris) in Newark, New Jersey (U.S.A.), were studied during summer (1978) and winter (1978–1979) months. Population densities of approximately 400 dogs per square mile and a sex ratio of 3 males: 1 female were found for each of three study areas. Free-ranging dogs were typically large, owned individuals; strays were a monority. Of four different group types identified in this study (familiar, unfamiliar, people-mediated and estrous), the first three are described. The social organization during non-breeding periods was characterized by a majority of solitary individuals. The relatively few groups observed rarely contained more than two individuals. Familiarity between dogs was the primary basis of sociality, although the distance a dog was from its home-site, its body size, age and ownership status also influenced social behavior. There was no seasonal variation in these patterns. Aggression was rare and mutual avoidance was the primary spacing mechanism. However, of those agonistic interactions that did occur, unfamiliar dogs were involved 5–15 times more frequently than familiar dogs.Home-range movements of individuals were largely restricted to areas around the home-site. Home-range size was correlated to a dog's ownership status much more than body size. There was no evidence of territoriality.Resources such as food and shelter influenced the social organization by concentrating individuals in areas where these were available. Major, predictable changes in available food had no effect on social behavior. Likewise, environmental parameters such as human presence and weather conditions generally influenced dog behavior indirectly.
Article
A group of three feral dogs (two males, one female) living in vacant buildings in St. Louis City was studied. They avoided close proximity with people and were active earlier and later in the day than the people and loose pets in the area. They found food while scavenging through human trash. The group's activities were usually initiated by the female of the group though otherwise there was no clear linear hierarchy and few ritualized displays of dominance or greeting. Specific “roles” within the group were observable. Though the female appeared to be pregnant during the study, puppies were never noted.
Article
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.
Article
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.
Article
Observations on the agonistic behaviour of 12 free-ranging dogs from two neighbouring groups were recorded in Katwa town, India. Both intra- and inter-group agonistic encounters were recorded on a seasonal basis. Mean (±S.E.) seasonal number of intra-group agonistic encounters of individual dog was greatest in winter (13.33±1.89) and then in late monsoon (12.33±1.99), when the females were lactating and in oestrus, respectively. Similarly, the mean (±S.E.) seasonal number of inter-group agonistic encounters of individual dog was greatest in winter (32.25±4.43) and then in late monsoon (27.75±2.01). There was a significant difference between the intra- and inter-group agonistic encounters. Dominance hierarchies were established among the adult dogs of either sex based on aggressive encounters. Although individual differences in agonism were observed, overall levels of aggression were higher among the adult females than for other groups. In contrast, overall levels of submission were higher among the juvenile males than for other groups. The results from this study suggest that reproductive season, sex and age have a significant effect on the agonistic behaviour of free-ranging dogs.
Article
The classic study of dog behavior gathered into one volume. Based on twenty years of research at the Jackson Laboratory, this is the single most important and comprehensive reference work on the behavior of dogs ever complied. "Genetics and the Social Behavior of the Dog is one of the most important texts on canine behavior published to date. Anyone interested in breeding, training, or canine behavior must own this book."—Wayne Hunthausen, D.V.M., Director of Animal Behavior Consultations "This pioneering research on dog behavioral genetics is a timeless classic for all serious students of ethology and canine behavior."—Dr. Michael Fox, Senior Advisor to the President, The Humane Society of the United States "A major authoritative work. . . . Immensely rewarding reading for anyone concerned with dog-breeding."—Times Literary Supplement "The last comprehensive study [of dog behavior] was concluded more than thirty years ago, when John Paul Scott and John L. Fuller published their seminal work Genetics and the Social Behavior of the Dog."—Mark Derr, The Atlantic Monthly "Genetics and the Social Behavior of the Dog is essential reading for anyone involved in the breeding of dogs. No breeder can afford to ignore the principles of proper socialization first discovered and articulated in this landmark study."-The Monks of New Skete, authors of How to Be Your Dog's Best Friend and the video series Raising Your Dog with the Monks of New Skete.
Article
In Experiment 1 male dogs were observed visiting either a tethered male or a tethered nonestrous female. The duration and frequency of visits and the parts of the stimulus animal that were investigated were recorded. In Experiment 2 male and female dogs were observed in a series of two-choice tests to determine their visiting preferences toward males and nonestrous and estrous females. In Experiment 3 similar two-choice olfactory tests were employed to demonstrate investigatory preferences for urine, feces, anal gland and vaginal secretions, saliva, and ear wax samples from the stimulus animals. The females were brought into artificial estrus with injections of estradiol benzoate, and changes were observed in their visiting and olfactory preferences and in their attractiveness to male dogs. Males spent less time visiting males than they spent visiting females, regardless of the physiological condition of the latter. However, the visiting time devoted to estrous females was longer than that spent with females not in heat. These preferences may be partially explained by the findings that male dogs find estrous urine and vaginal secretions more attractive than their nonestrous counterparts and that they prefer female urine and feces compared to corresponding samples from male dogs. Following the induction of artificial estrus, female subjects spent significantly more time investigating male dogs and male urine when compared to female dogs and female urine, respectively. Male dogs and to a lesser extent bitches urinated more frequently in the vicinity of the preferred animals and odor stimuli.
Article
The relationship between visual and olfactory cues and their relative importance in interactions between pairs of adult male ring-tailed lemurs was determined. Visual and olfactory contact was controlled. When visualcontact was prevented. little behavior occurred. When visual contact was permitted, olfactory cues altered behavior.
Article
Performed large bilateral lesions of the medial preoptic-anterior hypothalamic (MP-AH) continuum on 20 mixed breed adult male dogs in 2 experiments. These lesions, which eliminate copulatory behavior in male rats and cats, immediately abolished or greatly impaired the Ss' copulatory behavior. MP-AH lesions interfered with critical aspects of the mediation of sexual behavior aside from any alteration of gonadotrophin secretion or removal of forebrain androgen target tissue. The behavioral effects of the lesions could not be attributed to suppression of erectile or ejaculatory responses. The lesions also abolished or greatly impaired male urine-marking behavior, but did not influence aggressive behavior as observed in dominance-subordination relationships. (21/2 p ref)
Article
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.