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...h reinforcement, but suggested that it was brought about by pleiotropic effects associated with other changes evolved in isolation (see also Passmore 1985, Verrel & Arnold 1989, Dempster et al. 1993, =-=Andersson 1994-=-). On the other hand, the profound difference between male and female AGS within the same species (46%) suggests that AGS is used to signal sexual identity (see Schulte et al. 1995b, Rosell & Sun 1999...

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...signals indicate that the first cue (in this case chemical) functions to alert the receiver to the presence of the second cue (visual), increasing the probability of its detection 37 and recognition (=-=Endler 1992-=-, 1993, Wiley 1994). The corresponding results of the two types of aggressive responses measured (i.e. direct and overnight responses) indicate that discrimination of heterospecific scent marks is not...

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... excludes competitors from a specific resource or resources”. Territoriality may be expected to evolve when the benefits gained from exclusive access to limited resources exceed the costs of defence (=-=Brown 1964-=-, Stamps 1994). Costs of territoriality can be minimized if resident animals advertise their occupation of an area in order to deter intrusion and avoid escalated encounters with conspecifics. Adverti...

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...encounter (Gosling 1982, Gosling & Mckay 1990). It is therefore critical that signallers maintain their scent in such a way that maximizes the success of matching (Gosling 1986, Roberts & Lowen 1997, =-=Gosling & Roberts 2001-=-). This is achieved both by replenishing their own scent marks on a regular basis and by countermarking any scent deposited by competitors within their territory or area of dominance (Roberts 1998, Ri...

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...en argued that chemical signals are especially effective in this regard (Gosling 1986). If an animal has scent marked an area comprehensively, it must have inhabited it at least long enough to do so (=-=Gosling 1982-=-). Additionally, the signal remains active even when its author is absent from an area. Mammalian scent marking is often associated with territorial defence (e.g. Gosling 1990). It is widely accepted ...

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...e to identify the marker. Resident aardwolves (Proteles cristatus) sniffed neighbour’s marks significantly longer when found inside of their territories than at the borders (the ‘centre-edge effect’, =-=Falls 1982-=-, Sliwa & Richardson 1998). Further studies should clarify this issue for beavers. Species discrimination The results in paper VI confirm my hypothesis that Eurasian beavers discriminate between scent...

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...ritories, while the deposits of AGS sometimes found at latrines appear to have another function. The primary roles of skin glands of carnivores are the maintenance of the pelage and thermoregulation (=-=Gorman & Trowbridge 1989-=-). The same scent may also code for different information and thus serve multiple functions (e.g. Quay & Müller-Schwarze 1971, Epple et al. 1979, Johnston 1985), while several different scents may car...

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...Canis lupus) concentrated scent marks at the periphery of the territory. The same pattern was also found for the Eurasian beaver (this study, paper I & II), and for many other mammals (Aleksiuk 1968, =-=Kruuk 1978-=-, Kruuk et al. 1984, Smith et al. 1989, Richardson 1991, Sun et al. 1994, Gese & Ruff 1997, Sillero-Zubiri & Macdonald 1998, Brashares & Arcese 1999). In this manner, intruding beaver, upon entering a...

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...reat (Richardson 1991), a pattern of behaviour called countermarking (see also below). Scent countermarking is a common phenomenon among mammals and numerous functions have been proposed for it (e.g. =-=Ewer 1968-=-, Ralls 1971, Johnston et al. 1994, Wilcox & Johnston 1995, Roberts 1998, Sliwa & Richardson 1998, Ferkin 1999, Roberts & Dunbar 2000). In addition, overmarking and destroying a scent mound may mask i...

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...and reproductive status or mood, promoting synchronisation of reproductive cycles, attracting members of the opposite sex, labelling resources, and reassurance/confidence (see for example, reviews by =-=Eisenberg & Kleiman 1972-=-, Johnson 1973, Müller-Schwarze 1974, Thiessen & Rice 1976, Henry 1977, Brown 1979, Brown & Macdonald 1985, Kruuk 1992, Branch 1993, Lazaro-Perea et al. 1999). Scent marks therefore might serve severa...

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...e same plants. Thus, other factors than diet may be in part responsible for the observed difference (e.g. bacterial flora: Albone et al. 1977, Walro & Svendsen 1982, genetically based components: see =-=Halpin 1986-=-). The suggestion that a reduced aggressive response toward scent marks of the North American beaver is based on chemical differences between the two species is to a greater extent supported by AGS in...

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...is a common phenomenon among mammals and numerous functions have been proposed for it (e.g. Ewer 1968, Ralls 1971, Johnston et al. 1994, Wilcox & Johnston 1995, Roberts 1998, Sliwa & Richardson 1998, =-=Ferkin 1999-=-, Roberts & Dunbar 2000). In addition, overmarking and destroying a scent mound may mask information from other individuals. By covering a previously deposited scent with its own scent, an animal may ...

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...phenomenon One mechanism by which individuals may reduce defence costs is to reduce aggression towards familiar occupants of neighbouring territories, known as the dear enemy phenomenon (Fisher 1954, =-=Krebs 1982-=-, Ydenberg et al. 1988, Temeles 1994). Once territorial boundaries have been established, a territorial neighbour poses less threat to an individual’s territory and an aggressive response to its displ...

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...ular weight below 300. It could be advantageous for a swimming mammal such as the beaver to present chemical signals in the form of lipid substances that would concentrate at the air-water interface (=-=Albone 1984-=-). By lubricating the fur with AGS, which would be released into the water, beavers could also act as a ”living scent mark”. As AGS is insoluble in water (Svendsen 1978), beavers downstream would rece...

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...t is widely accepted that mammals scent mark their territories to advertise their occupancy and ownership of the territory (e.g. Peters & Mech 1975, Macdonald 1980, Erlinge et al. 1982, Gosling 1982, =-=Gorman & Mills 1984-=-, Smith et al. 1989, Sillero-Zubiri & Macdonald 1998), but it is still under debate how scent marks actually function in terms of territory maintenance (Gorman 1990, Gosling 1990, Richardson 1991, 199...

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.../excretions such as faeces and urine also may be used. Urine and faeces may be ideal substances for scent marking because they have a minimal energetic cost to the signaller (e.g. Gosling 1981, 1985, =-=Brashares & Arcese 1999-=-). Recent studies have demonstrated that scent types can carry different information and 18 thus have different functions (Johnston et al. 1993). For instance, the study by Gorman et al. (1978) on ott...

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... resources, and reassurance/confidence (see for example, reviews by Eisenberg & Kleiman 1972, Johnson 1973, Müller-Schwarze 1974, Thiessen & Rice 1976, Henry 1977, Brown 1979, Brown & Macdonald 1985, =-=Kruuk 1992-=-, Branch 1993, Lazaro-Perea et al. 1999). Scent marks therefore might serve several functions, which may change or vary with the time of year or the location of the mark. However, these alternative hy...

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...mbers of the opposite sex, labelling resources, and reassurance/confidence (see for example, reviews by Eisenberg & Kleiman 1972, Johnson 1973, Müller-Schwarze 1974, Thiessen & Rice 1976, Henry 1977, =-=Brown 1979-=-, Brown & Macdonald 1985, Kruuk 1992, Branch 1993, Lazaro-Perea et al. 1999). Scent marks therefore might serve several functions, which may change or vary with the time of year or the location of the...

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...ability to control the resources contained within defended areas (Parker 1974, Zahavi 1975). Although examples of visual and auditory signals functioning as territorial advertisement are common (e.g. =-=Hailman 1977-=-, Catchpole 1982), it has been argued that chemical signals are especially effective in this regard (Gosling 1986). If an animal has scent marked an area comprehensively, it must have inhabited it at ...

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...ts display would add unnecessary costs to territorial defence. Strangers, however, pose a greater threat and a heightened aggressive response might well be worth the cost of time and energy expended (=-=Jaeger 1981-=-, Temeles 1994). Other than increased visitation to ESMs marked with stranger castoreum, Schulte (1993, 1998) found little support for the dear enemy phenomenon in the North American beaver and conclu...

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...getic cost to the signaller (e.g. Gosling 1981, 1985, Brashares & Arcese 1999). Recent studies have demonstrated that scent types can carry different information and 18 thus have different functions (=-=Johnston et al. 1993-=-). For instance, the study by Gorman et al. (1978) on otters showed that deposits of spraints and urine might be used in the maintenance of otter territories, while the deposits of AGS sometimes found...

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...ttern was also found for the Eurasian beaver (this study, paper I & II), and for many other mammals (Aleksiuk 1968, Kruuk 1978, Kruuk et al. 1984, Smith et al. 1989, Richardson 1991, Sun et al. 1994, =-=Gese & Ruff 1997-=-, Sillero-Zubiri & Macdonald 1998, Brashares & Arcese 1999). In this manner, intruding beaver, upon entering a foreign territory, quickly discover that the area is already occupied. This general patte...

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...months, and both sexes defend their territories by scent marking (Wilsson 1971, Buech 1995). A variety of functions have also been assumed for scent marks in the beaver (see Dugmore 1914, Green 1936, =-=Aleksiuk 1968-=-, Butler & Butler 1979, MüllerSchwarze & Heckman 1980, Svendsen 1980a, Rosell & Bergan 1998). However, by testing alternative hypotheses, Houlihan (1989) confirmed the territorial function of North Am...

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... signalling, the main mechanism involved is thought to be “scent-matching”, in which competitors or mates match the odour from scent marks with the odour of conspecifics they encounter (Gosling 1982, =-=Gosling & Mckay 1990-=-). It is therefore critical that signallers maintain their scent in such a way that maximizes the success of matching (Gosling 1986, Roberts & Lowen 1997, Gosling & Roberts 2001). This is achieved bot...

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...iated dispersal in a downstream direction after ice-out. However, I do not know the main direction of dispersal in my study area, and beavers have been shown to disperse both upstream and downstream (=-=Leege 1968-=-, Van Deelen & Pletscher 1996). Another explanation for a predominance of upstream marking would be that intruders entering from a downstream direction automatically receive an almost continual flow o...

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...trol the resources contained within defended areas (Parker 1974, Zahavi 1975). Although examples of visual and auditory signals functioning as territorial advertisement are common (e.g. Hailman 1977, =-=Catchpole 1982-=-), it has been argued that chemical signals are especially effective in this regard (Gosling 1986). If an animal has scent marked an area comprehensively, it must have inhabited it at least long enoug...

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...rial defence (e.g. Gosling 1990). It is widely accepted that mammals scent mark their territories to advertise their occupancy and ownership of the territory (e.g. Peters & Mech 1975, Macdonald 1980, =-=Erlinge et al. 1982-=-, Gosling 1982, Gorman & Mills 1984, Smith et al. 1989, Sillero-Zubiri & Macdonald 1998), but it is still under debate how scent marks actually function in terms of territory maintenance (Gorman 1990,...

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...llenged, but that those that scent-marked at higher frequencies were subjected to fewer agonistic encounters than those marking at lower rates (i.e. the status advertisement hypothesis was supported (=-=Gosling 1990-=-)). However, many species place scent marks throughout their territories, sometimes at a higher density near more frequently used trails, dens, lodges, or sleeping sites (Müller-Schwarze 1983, Gosling...

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...ttracting members of the opposite sex, labelling resources, and reassurance/confidence (see for example, reviews by Eisenberg & Kleiman 1972, Johnson 1973, Müller-Schwarze 1974, Thiessen & Rice 1976, =-=Henry 1977-=-, Brown 1979, Brown & Macdonald 1985, Kruuk 1992, Branch 1993, Lazaro-Perea et al. 1999). Scent marks therefore might serve several functions, which may change or vary with the time of year or the loc...

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... concentrated scent marks at the periphery of the territory. The same pattern was also found for the Eurasian beaver (this study, paper I & II), and for many other mammals (Aleksiuk 1968, Kruuk 1978, =-=Kruuk et al. 1984-=-, Smith et al. 1989, Richardson 1991, Sun et al. 1994, Gese & Ruff 1997, Sillero-Zubiri & Macdonald 1998, Brashares & Arcese 1999). In this manner, intruding beaver, upon entering a foreign territory,...

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...e alternative hypotheses will not be the main focus here. For many years it was believed that scent marks help deter intruders from entering a territory, or at least to intimidate them (Hediger 1949, =-=Geist 1964-=-, Johnson 1973). The 15 intimidation hypothesis states that intruders may interpret scent marks as a threat with immediate physical attack if they are encountered by the resident (Hediger 1949, Richar...

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...99). A masking hypothesis has been proposed for many species that scent mark in situations that suggest territorial or home area defence and/or advertisement of dominance (Mertl 1977, Macdonald 1979, =-=Hurst 1987-=-, 1990). Johnston et al. (1994) suggested that in golden hamsters (Mesocricetus auratus) countermarking might have competitive functions, because after test males investigated the marks of two individ...

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...ate actions to take (i.e. signal detection theory, see Bradbury & Vehrencamp 1998). Thus, similar sniffing durations, or a lack of preference, does not indicate inability to discriminate (Brown 1979, =-=Johnston 1993-=-, Gouat et al. 1998), but can be interpreted as a process of decision-making. A similar behaviour has also been described for tree shrews (Tupaia belangeri) where the presentation of hetereospecific s...

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...icted to be most frequent when transient animals from other families are most likely to enter occupied areas, i.e. in spring or early summer when dispersal of 2-years-olds normally occurs (Beer 1955, =-=Bergerud & Miller 1977-=-, Molini et al. 1980, Svendsen 1980a). The North American beaver scent marks occur most often during May and June following birth and the dispersal of 2-year-olds (MüllerSchwarze & Heckman 1980, Svend...

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...nificant odours by mammals occurs in many organs that pass chemicals to the external environment. The major sources of odours used in territory defence are the skin glands (e.g. Müller-Schwarze 1983, =-=Flood 1985-=-), but metabolic byproducts/excretions such as faeces and urine also may be used. Urine and faeces may be ideal substances for scent marking because they have a minimal energetic cost to the signaller...

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...sling 1990). Social odours are a limited resource whether the animal use faeces, urine, or secretion from skin glands (sebaceous and sweat glands (apocrine and eccrine glands)) (Müller-Schwarze 1983, =-=Gorman 1984-=-a). Scent marking can also involve a significant investment in terms of time and energy (Gosling 1986) including the cost of reduced growth rate and body size (Gosling et al. 2000). Given these constr...

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...metabolic byproducts/excretions such as faeces and urine also may be used. Urine and faeces may be ideal substances for scent marking because they have a minimal energetic cost to the signaller (e.g. =-=Gosling 1981-=-, 1985, Brashares & Arcese 1999). Recent studies have demonstrated that scent types can carry different information and 18 thus have different functions (Johnston et al. 1993). For instance, the study...

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... However, these alternative hypotheses will not be the main focus here. For many years it was believed that scent marks help deter intruders from entering a territory, or at least to intimidate them (=-=Hediger 1949-=-, Geist 1964, Johnson 1973). The 15 intimidation hypothesis states that intruders may interpret scent marks as a threat with immediate physical attack if they are encountered by the resident (Hediger ...

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... from the underlying scent, thus making it difficult or impossible to perceive individual signatures in it. However, it is unlikely that countermarks will completely cover the competitor’s scent (see =-=Johnston et al. 1995-=-, Hurst & Rich 1999). A masking hypothesis has been proposed for many species that scent mark in situations that suggest territorial or home area defence and/or advertisement of dominance (Mertl 1977,...

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... opposite sex, labelling resources, and reassurance/confidence (see for example, reviews by Eisenberg & Kleiman 1972, Johnson 1973, Müller-Schwarze 1974, Thiessen & Rice 1976, Henry 1977, Brown 1979, =-=Brown & Macdonald 1985-=-, Kruuk 1992, Branch 1993, Lazaro-Perea et al. 1999). Scent marks therefore might serve several functions, which may change or vary with the time of year or the location of the mark. However, these al...

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...riality is a characteristic that is not adaptive and has not been selected for, but might evolve when two species compete for some material resource when they occur in the same habitat (see also e.g. =-=Catchpole 1978-=-, Greenberg et al. 1996, Griffis & Jaeger 1998). This implies that a territorial response toward heterospecific scent marks should be based on individual experiences only, and not on autonomically con...

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...nds)) (Müller-Schwarze 1983, Gorman 1984a). Scent marking can also involve a significant investment in terms of time and energy (Gosling 1986) including the cost of reduced growth rate and body size (=-=Gosling et al. 2000-=-). Given these constraints, scent marks should not be deployed at random, but instead in an organised pattern that maximises their chance of being discovered by the individuals to whom they are direct...

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... adaptive and has not been selected for, but might evolve when two species compete for some material resource when they occur in the same habitat (see also e.g. Catchpole 1978, Greenberg et al. 1996, =-=Griffis & Jaeger 1998-=-). This implies that a territorial response toward heterospecific scent marks should be based on individual experiences only, and not on autonomically controlled (Paquet 1991) or innate mechanisms. Th...

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...Rosell & Bergan 1998). However, by testing alternative hypotheses, Houlihan (1989) confirmed the territorial function of North American beaver scent marks and rejected other interpretations (see also =-=Hodgdon 1978-=-, Müller16 Schwarze & Heckman 1980, Svendsen 1980a, Houlihan 1989, Welsh & Müller-Schwarze 1989, Schulte 1998). To-date, only anecdotal observations exist for the functions of scent marking in territo...

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...efend an area, since only those successfully dominating the area can ensure that their marks both predominate (Gosling 1982) and are more recently deposited than those of any challenging competitors (=-=Hurst 1993-=-, Hurst & Rich 1999). The countermarking may therefore advertise that the territory is occupied and signal the costs of competition if the threat is ignored (e.g. Gosling 1990, Roberts & Dunbar 2000)....

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...cent, thus making it difficult or impossible to perceive individual signatures in it. However, it is unlikely that countermarks will completely cover the competitor’s scent (see Johnston et al. 1995, =-=Hurst & Rich 1999-=-). A masking hypothesis has been proposed for many species that scent mark in situations that suggest territorial or home area defence and/or advertisement of dominance (Mertl 1977, Macdonald 1979, Hu...

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... a pattern of behaviour called countermarking (see also below). Scent countermarking is a common phenomenon among mammals and numerous functions have been proposed for it (e.g. Ewer 1968, Ralls 1971, =-=Johnston et al. 1994-=-, Wilcox & Johnston 1995, Roberts 1998, Sliwa & Richardson 1998, Ferkin 1999, Roberts & Dunbar 2000). In addition, overmarking and destroying a scent mound may mask information from other individuals....

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...h sexes defend their territories by scent marking (Wilsson 1971, Buech 1995). A variety of functions have also been assumed for scent marks in the beaver (see Dugmore 1914, Green 1936, Aleksiuk 1968, =-=Butler & Butler 1979-=-, MüllerSchwarze & Heckman 1980, Svendsen 1980a, Rosell & Bergan 1998). However, by testing alternative hypotheses, Houlihan (1989) confirmed the territorial function of North American beaver scent ma...

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... covered in this manner. Indeed, swimming beavers keep their nostrils at the water level, thus enabling them to sense chemical messages from neighbouring beavers concentrated within the surface film (=-=Grønneberg & Lie 1984-=-) (paper II) (see also below). Organs (odorants) used in territorial defence My results in paper III supported the prediction that castoreum was most frequently deposited on scent marks (96 of 96) and...

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...s, Houlihan (1989) confirmed the territorial function of North American beaver scent marks and rejected other interpretations (see also Hodgdon 1978, Müller16 Schwarze & Heckman 1980, Svendsen 1980a, =-=Houlihan 1989-=-, Welsh & Müller-Schwarze 1989, Schulte 1998). To-date, only anecdotal observations exist for the functions of scent marking in territorial defence by Eurasian beavers. Studies of scent marking in the...

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...sis) predicts that animals should mark where neighbours are most likely to encounter marks (Gosling 1986, Smith et al. 1989), and preferentially along borders adjacent to the most threatening rivals (=-=Johansson & Liberg 1996-=-). If scent-marking activity is correlated with population density (highly challenged), a positive correlation between number of neighbouring territories (or number of neighbouring individuals) and nu...

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...r, 2n=48) as a model to further elucidate this topic. Both the Eurasian and the North American beavers (C. canadensis, 2n=40) are strongly territorial and aggressive encounters are not uncommon (e.g. =-=Lavrov & Orlov 1973-=-, Piechocki 1977, Svendsen 1989, Nolet & Rosell 1994). Beavers usually deposit scent (castoreum and anal gland secretion (AGS), see below) onto small piles of mud and debris close to the water's edge ...

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...ce/confidence (see for example, reviews by Eisenberg & Kleiman 1972, Johnson 1973, Müller-Schwarze 1974, Thiessen & Rice 1976, Henry 1977, Brown 1979, Brown & Macdonald 1985, Kruuk 1992, Branch 1993, =-=Lazaro-Perea et al. 1999-=-). Scent marks therefore might serve several functions, which may change or vary with the time of year or the location of the mark. However, these alternative hypotheses will not be the main focus her...

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...ing is predicted to be most frequent when transient animals from other families are most likely to enter occupied areas, i.e. in spring or early summer when dispersal of 2-years-olds normally occurs (=-=Beer 1955-=-, Bergerud & Miller 1977, Molini et al. 1980, Svendsen 1980a). The North American beaver scent marks occur most often during May and June following birth and the dispersal of 2-year-olds (MüllerSchwar...

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...lled by them (e.g. Rosell & Czech, 1999). Dickman & Doncaster (1984) suggested that similar chemicals eliciting avoidance in rodents may commonly occur in the faeces and urine of carnivores (see also =-=Bininda-Emonds et al. 2001-=-). This is supported by observations that rodents often avoid the odours of carnivores with which there has been no evolutionary contact (Stoddart 1982a,b, Nolte et al. 1994, see also Roberts et al. 2...

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...entially and display a better memory for the odour of the top-scent donor than that of the bottom-scent donor (Johnston et al. 1994, 1995, 1997a,b, Wilcox & Johnston 1995, 33 Johnston & Bhorade 1998, =-=Ferkin et al. 1999-=-, Woodward et al. 1999). This preference for the top scent suggests that these animals treat the odour of the top-scent donor as being more important or having greater value than that of the bottom-sc...

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...“dear enemy” phenomenon One mechanism by which individuals may reduce defence costs is to reduce aggression towards familiar occupants of neighbouring territories, known as the dear enemy phenomenon (=-=Fisher 1954-=-, Krebs 1982, Ydenberg et al. 1988, Temeles 1994). Once territorial boundaries have been established, a territorial neighbour poses less threat to an individual’s territory and an aggressive response ...

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... visual and auditory signals functioning as territorial advertisement are common (e.g. Hailman 1977, Catchpole 1982), it has been argued that chemical signals are especially effective in this regard (=-=Gosling 1986-=-). If an animal has scent marked an area comprehensively, it must have inhabited it at least long enough to do so (Gosling 1982). Additionally, the signal remains active even when its author is absent...

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...ed along with the speciation process. Also, this is of particular interest in the wake of introductions of the North American beaver to Eurasia and the impending range concurrence of the two species (=-=Lahti 1995-=-). Main aims of the study I hypothesize that scent marking plays an important role in territory defence of free-ranging Eurasian beavers (Figure 1). Based on the main issues outlined above, I investig...

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...Nordiska ministerrådet 1984 for comparison) and probably forage on many of the same plants. Thus, other factors than diet may be in part responsible for the observed difference (e.g. bacterial flora: =-=Albone et al. 1977-=-, Walro & Svendsen 1982, genetically based components: see Halpin 1986). The suggestion that a reduced aggressive response toward scent marks of the North American beaver is based on chemical differen...

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...le conspecifics with their own odours. That is, they responded in a way similar to the over-marking shown by many other species (e.g. Hurst 1987, 1990, 1993, Johnston et al. 1994, 1995, Roberts 1998, =-=Bel et al. 1999-=-, Ferkin 1999). Also, the lack of a response to ESMs without castoreum indicated that beavers were responding to the smell of castoreum and not to the sight of the scent mound. Studies of North Americ...

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...hanisms evolved through reinforcement, but suggested that it was brought about by pleiotropic effects associated with other changes evolved in isolation (see also Passmore 1985, Verrel & Arnold 1989, =-=Dempster et al. 1993-=-, Andersson 1994). On the other hand, the profound difference between male and female AGS within the same species (46%) suggests that AGS is used to signal sexual identity (see Schulte et al. 1995b, R...

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...ng conspecifics (e.g. Roper et al. 1993, Gosling & Wright 1994, Ramsay & Giller 1996), but few studies have examined the prevalence of countermarking between heterospecifics (see however Paquet 1991, =-=Fornasieri & Roeder 1992-=-). Interspecific territoriality might evolve when species with overlapping ecological requirements interact (Simmons 1951). The greater the degree of overlap between species, the greater the competiti...

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...sling 1990). Social odours are a limited resource whether the animal use faeces, urine, or secretion from skin glands (sebaceous and sweat glands (apocrine and eccrine glands)) (Müller-Schwarze 1983, =-=Gorman 1984-=-a). Scent marking can also involve a significant investment in terms of time and energy (Gosling 1986) including the cost of reduced growth rate and body size (Gosling et al. 2000). Given these constr...

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...se inhibited further challenges. Countermarking also ensures that own scent marks always remain the most recently deposited. Such behaviour is readily seen among conspecifics (e.g. Roper et al. 1993, =-=Gosling & Wright 1994-=-, Ramsay & Giller 1996), but few studies have examined the prevalence of countermarking between heterospecifics (see however Paquet 1991, Fornasieri & Roeder 1992). Interspecific territoriality might ...

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...racteristic that is not adaptive and has not been selected for, but might evolve when two species compete for some material resource when they occur in the same habitat (see also e.g. Catchpole 1978, =-=Greenberg et al. 1996-=-, Griffis & Jaeger 1998). This implies that a territorial response toward heterospecific scent marks should be based on individual experiences only, and not on autonomically controlled (Paquet 1991) o...

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...Müller-Schwarze 1974, Brown 1979, Müller-Schwarze 1983, Feoktistova 1995). The ability to discriminate odours from different individuals has been documented for several mammalian species (reviewed in =-=Halpin 1980-=-, 1986). However, whether the Eurasian beaver can recognise an intruder (i.e. is this a potential intruder?) and discriminate a neighbour from a stranger or a conspecific from a heterospecific (i.e. w...

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...been described for tree shrews (Tupaia belangeri) where the presentation of hetereospecific scent marks elicited intense olfactory investigation, but no equivalent increase in scent marking activity (=-=Holst & Buergel-Goodwin 1975-=-). If the chemical signal present in castoreum and AGS of each species to some extent matches the chemical template of the other species, this might have led to the undifferentiated sniffing duration ...

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... mood, promoting synchronisation of reproductive cycles, attracting members of the opposite sex, labelling resources, and reassurance/confidence (see for example, reviews by Eisenberg & Kleiman 1972, =-=Johnson 1973-=-, Müller-Schwarze 1974, Thiessen & Rice 1976, Henry 1977, Brown 1979, Brown & Macdonald 1985, Kruuk 1992, Branch 1993, Lazaro-Perea et al. 1999). Scent marks therefore might serve several functions, w...

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...n order to deposit the exudates (Wilsson 1971, Svendsen 1978). It is suspected that castoreum is the most frequently used of the two during the scent-marking of territories (e.g. Schulte et al. 1994, =-=Bergan 1996-=-). Castoreum may be an ideal substance for scent marking the territory because it has a minimal energetic cost to the signaller. Selection for effective signal-sending behaviour harnesses odours that ...

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...d and debris close to the water's edge (e.g. Wilsson 1971, Svendsen 1980a). All age-classes, except kits younger than 5 months, and both sexes defend their territories by scent marking (Wilsson 1971, =-=Buech 1995-=-). A variety of functions have also been assumed for scent marks in the beaver (see Dugmore 1914, Green 1936, Aleksiuk 1968, Butler & Butler 1979, MüllerSchwarze & Heckman 1980, Svendsen 1980a, Rosell...

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... et al. 1982, Gosling 1982, Gorman & Mills 1984, Smith et al. 1989, Sillero-Zubiri & Macdonald 1998), but it is still under debate how scent marks actually function in terms of territory maintenance (=-=Gorman 1990-=-, Gosling 1990, Richardson 1991, 1993). Alternative hypotheses, however, have been proposed for scent marking in mammals: identification of species, subspecies, group, or individuals, signalling socia...

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...nger than 5 months, and both sexes defend their territories by scent marking (Wilsson 1971, Buech 1995). A variety of functions have also been assumed for scent marks in the beaver (see Dugmore 1914, =-=Green 1936-=-, Aleksiuk 1968, Butler & Butler 1979, MüllerSchwarze & Heckman 1980, Svendsen 1980a, Rosell & Bergan 1998). However, by testing alternative hypotheses, Houlihan (1989) confirmed the territorial funct...

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... pelage and thermoregulation (Gorman & Trowbridge 1989). The same scent may also code for different information and thus serve multiple functions (e.g. Quay & Müller-Schwarze 1971, Epple et al. 1979, =-=Johnston 1985-=-), while several different scents may carry the same information (Baldwin & Meese 1977, Roeder 1980, Martin & Beauchamp 1982). Beaver possess two pairs of scent producing organs, castor sacs and anal ...

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...marks is essential for the animal if it is to identify potential mates, competitors, and territory owners (Johnston et al. 1995, 1997a,b, Wilcox & Johnston 1995, Johnston & Bhorade 1998, Ferkin 1999, =-=Kohli & Ferkin 1999-=-). Johnston et al. (1994) outlined three hypotheses to explain what happens when scent marks of two conspecifics overlap. The first hypothesis, called scent-blending, states that the two scents will m...

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...torial defence by Eurasian beavers. Studies of scent marking in the Eurasian beaver typically have focused on the behaviour of only a few animals or of captive/semi captive individuals (Wilsson 1971, =-=Anderson & Westerling 1984-=-, Nolet & Rosell 1994). Understanding the functions of scent marking in Eurasian beaver territorial defence may contribute important findings for a better understanding of this species’ communication ...

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...o code for different information and thus serve multiple functions (e.g. Quay & Müller-Schwarze 1971, Epple et al. 1979, Johnston 1985), while several different scents may carry the same information (=-=Baldwin & Meese 1977-=-, Roeder 1980, Martin & Beauchamp 1982). Beaver possess two pairs of scent producing organs, castor sacs and anal glands (Svendsen 1978, Walro & Svendsen 1982, Valeur 1988), and both are suspected to ...

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...e main purpose of a beaver’s investigation of an ESM is to identify the sender, and then, based on the information obtained, decide what appropriate actions to take (i.e. signal detection theory, see =-=Bradbury & Vehrencamp 1998-=-). Thus, similar sniffing durations, or a lack of preference, does not indicate inability to discriminate (Brown 1979, Johnston 1993, Gouat et al. 1998), but can be interpreted as a process of decisio...

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...and reassurance/confidence (see for example, reviews by Eisenberg & Kleiman 1972, Johnson 1973, Müller-Schwarze 1974, Thiessen & Rice 1976, Henry 1977, Brown 1979, Brown & Macdonald 1985, Kruuk 1992, =-=Branch 1993-=-, Lazaro-Perea et al. 1999). Scent marks therefore might serve several functions, which may change or vary with the time of year or the location of the mark. However, these alternative hypotheses will...

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...4) remove or replace marks of others (Gosling 1982, 1985, 1986, Gorman 1984a). The scentmatching hypothesis has received support by studies of scent marking in several species, e.g. ferret (M. furo) (=-=Clapperton et al. 1988-=-), house mice (Mus domesticus) (Gosling & McKay 1990), suni antelope (Neotragus moschatus) (Somers et al. 1990), yellow mongoose (Cynictus penicillata) (Wenhold & Rasa 1994), and North American beaver...

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...xcept kits younger than 5 months, and both sexes defend their territories by scent marking (Wilsson 1971, Buech 1995). A variety of functions have also been assumed for scent marks in the beaver (see =-=Dugmore 1914-=-, Green 1936, Aleksiuk 1968, Butler & Butler 1979, MüllerSchwarze & Heckman 1980, Svendsen 1980a, Rosell & Bergan 1998). However, by testing alternative hypotheses, Houlihan (1989) confirmed the terri...

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...(paper II). The recently discovered vomeronasal organ in Eurasian beavers may play a significant role at the air-water interface but its importance for chemical communication in beavers is not known (=-=Døving et al. 1993-=-, Rosell & Pedersen 1999). However, the design of the beaver’s nose enables this amphibious animal to sample the chemical composition of its environment. Above water the beaver can inhale air and expo...

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... maintenance of the pelage and thermoregulation (Gorman & Trowbridge 1989). The same scent may also code for different information and thus serve multiple functions (e.g. Quay & Müller-Schwarze 1971, =-=Epple et al. 1979-=-, Johnston 1985), while several different scents may carry the same information (Baldwin & Meese 1977, Roeder 1980, Martin & Beauchamp 1982). Beaver possess two pairs of scent producing organs, castor...

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... recorded from a neighbour are broadcast to a resident from the territory boundary opposite the shared boundary, the residents treat neighbours and strangers equally aggressively (Wiley & Wiley 1977, =-=Falls 1978-=-, Trivers 1985). Therefore, animals living on adjacent territories should show a clearer dear enemy 35 phenomenon than animals on territories with undefended space between. Caley & Boutin (1987) found...

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...or a wealth of information including species, subspecies, social group, individuality, sex, age, social status and reproductive condition (e.g. Müller-Schwarze 1974, Brown 1979, Müller-Schwarze 1983, =-=Feoktistova 1995-=-). The ability to discriminate odours from different individuals has been documented for several mammalian species (reviewed in Halpin 1980, 1986). However, whether the Eurasian beaver can recognise a...

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...were individually marked (ear tagged and implanted with microchips). All beavers (both shot and live-trapped) were sexed (Rosell & Sun 1999), weighed and assigned to age classes based on body weight (=-=Hartman 1992-=-, Rosell & Pedersen 1999, Parker et al. 2001) (papers III-VI). However, in paper III the age of dead beavers was determined by examining tooth root closure and cementum annuli layers of the first mola...

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.... Distinguishing among multiple scent marks is essential for the animal if it is to identify potential mates, competitors, and territory owners (Johnston et al. 1995, 1997a,b, Wilcox & Johnston 1995, =-=Johnston & Bhorade 1998-=-, Ferkin 1999, Kohli & Ferkin 1999). Johnston et al. (1994) outlined three hypotheses to explain what happens when scent marks of two conspecifics overlap. The first hypothesis, called scent-blending,...