Scent-Marking in the Eurasian Beaver (Castor fiber) as a Means of Territory Defense

Beaver (Castor spp.) normally scent-mark by depositing castoreum and/or anal gland secretion on scent mounds close to the water's edge. The aim of this study was to investigate the hypothesis that the Eurasian beaver (C. fiber) scent-marks as a means of territory defense. Scent-marking behavior was studied during an entire year (April 1, 1995–March 31, 1996) in seven adjacent territories along 9.2 km of the B River in Telemark County, Norway. The number and location of fresh scent marks were recorded biweekly. The main results showed that: (1) the number of scent marks in territories was significantly higher in spring, when dispersal of subadults normally occurs than during the rest of the year; (2) the number of scent marks was clumped near territorial borders; and (3) the number of scent marks was significantly greater upstream than downstream of the lodge.     

Do Eurasian beavers smear their pelage with castoreum and anal gland secretion?

The scent-matching hypothesis postulates that scent marks provide an olfactory link between a resident owner and his territory, and that this enables intruding animals to recognize the chance of escalated conflicts. However, it is unclear if Eurasian beavers (Castor fiber) mark their own pelage with castoreum from their castor sacs (i.e., the same material used in territorial marking); and/or if beavers mark their pelage with anal gland secretion (AGS) from the anal glands to waterproof the pelage and to act as a living-scent mark. Chemical analysis (gas chromatography and mass spectrometry) of hair samples from 22 live-trapped beavers revealed that castoreum compounds were not present in any samples, AGS compounds were found from 3 animals (13.6%) around the cloaca, and the compound squalene was found in all the samples. Beavers may release castoreum directly into the water when it meets an intruder. Thereby, the scent mark in the water can provide an olfactory link between a resident owner and his territory. Squalene, in contrast to AGS, may be essential for keeping beaver pelts water-repellant.     

Castor sacs and anal glands of the north american beaver (Castor canadensis): their histology,development, and relationship to scent communication

Both sexes of beavers possess a pair of castor sacs and a pair of anal glands located in paired subcutaneous cavities between the pelvis and the base of the tail. The castor sacs are not glandular in the histological sense, hence references to these structures as preputial glands or castor glands are misnomers. The wall of the castor sacs is plicate and comprised of three distinct zones: an outer layer of vascular connective tissue, a two-to five-cell-thick layer of mitotic epithelial cells, and several densely packed layers of cornified epithelium which grade into more widely separated sheets toward the lumen. Monocultures of a gram-positive facultatively anaerobic bacterium were present in the lumen of all castor sac preparations. Differences in the frequency of castoreum deposition were not attributable to differences in the structure of the castor sacs. The anal glands of beavers are holocrine sebaceous glands. These glands develop more rapidly than the castor sacs. Anal gland tissue from embryos exhibited cellular characteristics associated with the production of sebum. Secretory activity was evident in all preparations. The relationship of castoreum and anal gland secretion to scent communication among beavers is discussed.     

Sexual Dimorphism in Territorial Scent Marking by Adult Eurasian Beavers (Castor fiber)

Mammalian scent marking is often associated with territorial defense. However, males and females may demonstrate different activity patterns and play different roles. Female mammals nurture the young during lactation, while males purportedly perform other tasks more frequently, such as territorial maintenance and defense. This paper investigates the contribution made by mated pairs of adult males and females to territorial scent-marking in the obligate monogamous Eurasian beavers (Castor fiber). We hypothesized that both sexes should show territorial behavior, and predicted that they deposit a higher proportion of scent marks at borders. We also hypothesized that a sexual dimorphism exists due to reproductive constraints on the females, and predicted that males should invest significantly more in scent-marking behavior than females during summer. We obtained behavioral data by radio tracking six mated pairs of Eurasian beavers during spring and summer 2000–2001 on two rivers in southeastern Norway. Our results showed that both males and females clustered their scent marks near territorial borders, but males deposited a larger number of scent marks than females and spent more time at borders. Males were also found to have a higher scent marking rate and scent marks per night than females during summer, but not during spring. Overall, scent marks per night were higher in males than females. We conclude that both males and females Eurasian beavers carry out territorial behavior by scent marking, but males carry a larger part of the territorial defense during summer when females lactate. Our results are discussed in the light of the codefense hypothesis.     

Beaver (Castor canadensis) responses to major phenolic and neutral compounds in castoreum

North American beaver (Castor canadensis) mark their territories with castoreum, a chemically complex secretion from their castor sacs. The phenolic and neutral fractions of castoreum have been shown to elicit specific behavioral responses from beavers in a field setting. Our objective was to identify compounds/mixtures that evoked responses similar to those stimulated by castoreum. We assayed recently identified phenolic compounds, some phenolics that had been determined to be biologically active in previous studies, the neutral compound borneol, and combinations of phenolic compounds, neutral compounds, and the two combined. Biological activity was measured by the elicitation and extent of specific responses and their strength (duration, frequency, and proportion of beavers responding). Generally, single compounds stimulated fewer responses than mixtures. A 26-compound mixture of phenolic and neutral compounds elicited responses in a similar proportion of trials as castoreum. However, responses to castoreum were stronger than to any synthetic sample. Further investigation of different measures of response, namely, elicitation, completeness, and strength, are deemed necessary to fully decipher the design of social odors.     

Bacterial contents of the anal and castor glands of beaver (Castor canadensis)

Bacterial contents of both the anal gland and castor gland of the beaver (Castor canadensis) were determined. Using our culture methods, no bacteria were isolated from the castor glands, but the anal gland contained high numbers of the aerobeEscherichia coli and the anaerobeBacteroides fragilis. The latter may be represented by several variants but facilities were not available for advanced anaerobic analysis. The relative numbers of each bacterial group and the group present were constant regardless of sex, age class, or colony of beaver. The bacterial fermentation hypothesis is rejected for castor gland section but remains possible for anal gland secretions based on variations seen inB. fragilis. The role of the products of both the castor gland and anal gland are discussed in relationship to scent communication in beaver.     

Experimental habitat scenting inhibits colonization by beaver,Castor canadensis

Unoccupied beaver (Castor canadensis) sites in New York State were for two years experimentally scented with a mixture of beaver castoreum and anal gland secretion. These sites were colonized less often than unscented control sites. The beaver is the first mammal to have been shown experimentally to use intraspecific odor cues when settling in vacant habitat. Territorial pheromones may be useful as repellents for beaver or other rodents.     

Neutral compounds from male castoreum of North American beaver,Castor canadensis

North American beavers (Castor canadensis) mark their territories with castoreum, the strong-smelling paste in their castor sacs. In their own territories, beavers respond with scent marking to experimental scent marks that consist of strange castoreum (or selected components). In part, the unique odor of castoreum is due to large amounts of phenolic compounds and neutral compounds. Purified neutral compounds were analyzed by GC. GC-MS, and NMR; identities of the neutral compounds were confirmed by comparing the properties of authentic compounds with those of the isolated compounds. We identified 13 neutral compounds that had not been reported before for castoreum. Most of these are oxygen-containing monoterpenes. Of the nine neutral compounds reported by Lederer (1949), only three are confirmed in our analysis; the other six neutral compounds are either absent or are not volatile enough to be detected by our methods. Eight compounds—6-methyl-l-heptanol, 4,6-dimethyl-l-heptanol, isopinocamphone, pinocamphone, two linalool oxides, and their acetates—were synthesized for structure identification and bioassays.     

Pheromonal activity of single castoreum constituents in beaver,Castor canadensis

Behavioral activity of single components of beaver castoreum was demonstrated for the first time. In four experiments samples were presented to free-ranging beaver in their family territories. First, responses to whole castoreum and anal gland secretion (AGS) from males and females were tested. Second, 24 compounds, known to be constituents of beaver castoreum, were individually screened for activity. Four of these consistently released immediate responses during the observation periods. These are the phenols 4-ethylphenol and 1,2-dihydroxybenzene and the ketones acetophenone and 3-hydroxyacetophenone. In the most complete responses, the beaver sniffed from the water, were attracted to the odor, swam toward its source, went on land, and then approached, sniffed, pawed, and scent-marked the artificial scent mound. 4-Ethoxyphenol, a compound not yet found in castoreum, also released these responses. Five additional compounds resulted in a few delayed visits to the samples during the night following the observations, as evidenced by destroyed scent mounds. These are 4-methyl-1,2-dihydroxybenzene, 4-methoxyacetophenone, 5-methoxysalicylic acid, salicylaldehyde, and 3-hydroxybenzoic acid. Third, mixtures of 24 and six compounds were tested. Responses to these mixtures could be as strong as those to whole castoreum. Fourth, the four regularly active compounds were tested in two additional beaver populations and proved to be active there, too. The response was strongest in the densest beaver population.     

Phenolic compounds from male castoreum of the North American beaver,Castor canadensis

North American beaver (Castor canadensis) mark their territories with castoreum, the contents of their castor sacs. In their territories, beaver respond with scent marking to experimental scent marks consisting of castoreum, or selected single components. In part, the unique odor of castoreum is due to large amounts of phenolic compounds. Purified phenolic components were analyzed by GC, GC-MS, and NMR; identifications were confirmed by comparing the spectra of synthetic phenols with those of the isolated phenols. Of the 15 phenols reported elsewhere, only five were confirmed in our analysis; the other 10 phenolic compounds are either absent or are not volatile enough to be detected by our methods. In addition, 10 phenolic compounds have been identified in this study that were not reported in the previous papers concerning the constitution of castoreum.     

Patterns of scent-mounding in a population of beaver (Castor canadensis)

The frequency and pattern of distribution of scent-mound construction were studied in a population of beaver in southeast Ohio from 1975 through 1977. The study addressed the questions of whether or not the frequency of scent-mound construction varied with season, site, year, and degree of contact with other family groups, and whether the pattern of scent-mounding activity was parsimonious with the idea of territorial marking. The number of scent-mounds constructed was determined weekly for each site throughout the ice-free season. Scent-mounding activity was highest in spring and declined and remained low during summer and fall. Significant differences were found amoung sites and over years. Contact with other resident family groups altered both frequency and pattern of scent-mound construction. Scent-mounds did not conform to a scent-fence model. The most parsimonious interpretation of function of odor cues deposited on scent-mounds is the effect on the motivational state of residents and nonresidents, increasing the confidence and reducing anxiety in residents smelling their own scent-mound and decreasing the confidence and increasing the readiness to flee in trespassers encountering a strange scent-mound.     

Seasonal variation of three ketones in the interdigital gland secretion of reindeer (Rangifer tarandus L.)

The seasonal variation of the three ketones, 6-methyl-2-heptanone, 7-methyl-3-octanone, and 7-methyl-1-octen-3-one, in the interdigital gland secretion of reindeer (Rangifer tarandus L.) has been investigated. A group of four adult animals, three females and one male, was used as a test group during a period of two years. Even though the three compounds are monitored in only a small number of test animals, it is evident that the ratios of the secretion's components vary over time. The variations are cyclic and are repeated from year to year.     

Red maple (Acer rubrum) inhibits feeding by beaver (Castor canadensis)

At many beaver (Castor canadensis) sites at Allegany State Park in New York State, red maple (Acer rubrum) is the only or one of the few tree species left standing at the ponds' edges. The relative palatability of red maple (RM) was studied in three ways. (1) At seven beaver sites, the available and utilized trees were recorded and an electivity index (E) computed. Of 15 tree species, RM ranked second or fourth lowest. (2) In experiment I, RM, sugar maple (A. saccharum, SM), and quaking aspen (Populus tremuloides) logs were presented cafeteria style at 10 colonies. RM was the least preferred. (3) Bark of RM was extracted with solvents. Aspen logs were painted (experiment II) or soaked (experiment III) with this RM extract and presented to beaver cafeteria-style, along with aspen and RM controls. This treatment rendered aspen logs less palatable, indicating that a chemical factor had been transferred.     

Effect of volatiles collected above fecal pellets on behavior of the rabbit,Oryctolagus cuniculus,tested in an experimental chamber

The fecal pellets of rabbits,Oryctolagus cuniculus, are coated with secretion from the anal gland, the odor from which functions as a territorial marker. The total volatiles collected from above fecal pellets and three different fractions prepared by washing the volatiles through either distilled water, 1 M sodium hydroxide, or 1 M hydrochloric acid were tested for their effect on the territorial confidence of individual animals. Tests were based on the neutral pen paradigm used in previous studies but employed a specially designed test chamber, the internal odor environment of which could be fully controlled. In the four series of bioassays a total of 140 separate tests were performed and 64 adult male rabbits were used. The results showed that the presence of the total volatiles influences the outcome of territorial competitions in pairs of male rabbits in favor of the individual from which they were derived. They were comparable with those obtained in earlier studies when fecal pellets were presented in the test pen. The effects of the different washing reagents on the total odor are illustrated by gas chromatograms, and the components most likely to be removed by the different treatments are suggested. The volatiles remaining after passing through distilled water had less effect on the rabbit's behavior than the total odor. Since the washing process not only removed water-soluble compounds but also led to a reduction in the concentrations of water-insoluble components introduced into the test chamber, either of these factors could have been responsible for the weakened effect. Fractions of the odor obtained by washing with either sodium hydroxide or hydrochloric acid did not influence the results of the tests. This suggests that the essential components of the olfactory signal include both basic and acidic compounds.     

Thietanes and dithiolanes from the anal gland of the stoat (Mustela erminea)

2-Propylthietane (1), 2-pentylthietane (3), and 3-propyl-1, 2-dithiolane (4) were identified as the sulfur-containing components of the male stoat anal gland secretion. Additional sulfur containing compounds identified in female stoat anal gland secretions were 2-ethylthietane (7) and 3-ethyl-1,2-dithiolane (8).     

Volatile ketones from the preorbital gland of reindeer (Rangifer t. tarandus L.)

The identification of 4-heptanone and 2-methyl-4-heptanone from the preorbital gland of reindeer (Rangifer t. tarandus L.) is reported. The secretion of the gland exhibit marked seasonal and sexual differences.     

Linalool in mandibular gland secretion ofColletes bees (Hymenoptera: Apoidea)

Uinalool was identified as the main component of the mandibular gland secretion in females and males of four species ofColletes bees, viz.,C. cunicularius, C. daviesanus, C. impunctatus andC. succinctus. It was also shown to be the dominant volatile compound in the same gland in maleC. floralis (the female of this species has not yet been investigated). Further, in another species,C. similis, linalool is present in the mandibular gland secretions of females and males, but the dominant volatile component in these secretions is geranial, together with neral. Females of the speciesC. fodiens seem to lack monoterpenes altogether; nonadecane is the dominant volatile compound of the cephalic secretion (based on analysis of a single individual). When linalool is put out in the area of nest aggregation ofC. cunicularius, where the males are patrolling, a distinct increase in flight activity is noted. The function of linalool is discussed on the basis of field observations.This report forms part XXI in the series Studies on Natural Odoriferous Compounds.     

Effect of volatiles collected above fecal pellets on behavior of the rabbit,Oryctolagus cuniculus,tested in an experimental chamber

Gas chromatographic techniques were used to trap and fractionate the volatile substances emanating from the fecal pellets of wild rabbits,Oryctolagus cuniculus, which incorporate the odor of the anal gland secretion. The volatiles were bubbled through water, trapped on Chromosorb 105, transferred to Silicone SF96 traps, passed through a GC capillary column, and retrapped on SF96. The behavioral effect of trapped and fractionated volatiles was determined in four series of bioassays involving 51 adult, male, wild-type rabbits in 112, ten-minute tests. The bioassay was based on the demonstration of territorial confidence by the rabbits under the influence of their own odor. In the bioassays of the unfractionated volatiles eluted from Chromosorb 105 and SF96 traps the animals were significantly more confident in the presence of the volatiles from their own fecal pellets. One of the two fractions of the total volatiles was ineffective while the effect of the other was less than that of the unfractionated odor. The results demonstrate that the gas chromatographic techniques can be used to manipulate complex mammalian odors with precision. Combined with a discriminative bioassay, this opens up the possibility of identifying the specific combinations of volatile substances involved in the formation of olfactory signals.     

Anal gland secretion of the ferret (Mustela putorius formaFuro)

2,2-Dimethylthietane(1),trans-2,3-dimethylthietane(2),cis-2,3-dimethylthietane(3), 2-propylthietane(4), 3,3-dimethyl-1,2-dithiolane(5),trans-3,4-dimethyl-1,2-dithiolane(6),cis-3,4-dimethyl-1,2-dithiolane(7), 2-pentylthietane (8), 3-propyl-1,2-dithiolane(9), quinoline(10), and indole(11) have been identified as components of the anal gland secretion of the ferret,Mustela putorius formafuro. Male and female ferret secretions were indistinguishable except during the breeding season.     

Bioactivity of beaver castoreum constituents using principal components analysis

North American beaver (Castor canadensis) were observed to sniff from the water and make land visits to some synthetic chemical components of castoreum placed on experimental scent mounds (ESM). In previous analysis, the elicitation (presence/absence), completeness, and/or strength (number, duration) of these key responses served as separate measures of biological activity. In this paper, we used principal components analysis (PCA) to combine linearly six related measures of observed response and one index of overnight visitation calculated over all trials. The first principal component accounted for a majority of the variation and allowed ranking of the samples based on their composite bioactivity. A second PCA, based only on response trials (excluding trials with no responses), showed that responses to the synthetic samples, once elicited, did not vary greatly in completeness or strength. None of the samples evoked responses as complete or strong as the castoreum control. Castoreum also elicited more multiple land visits (repeated visits to the ESM by the same individual or by more than one family member) than the synthetic samples, indicating that an understanding of the castoreum chemosignal requires consideration of responses by the family unit, and not just the land visit by the initial responder.