Some communicatory functions of scent marking in the cotton-top tamarin (Saguinus oedipus oedipus)

The ability of cotton-top tamarins to discriminate between scents from conspecifics and those from other tamarin species, and between scents from conspecific individuals was tested. Cotton-tops scent mark with specialized skin glands in the circumgenital area. Females possess larger glands than males and show more scent-marking behavior. In the first experiment, subjects were presented with a glass rod scented with either material collected from the surface of the scent glands of a conspecific female, with scent material from a female of a related species, or with an unscented rod. Glass rods carrying scent from conspecifics were sniffed more frequently than rods carrying scent from related tamarin species or unscented rods. A second experiment offered a choice between two glass rods, one scent marked by aSaguinus o. oedipus female, the second one scent marked by aSaguinus fusdcollis female. Shelves carrying rods that had been scent marked by conspecifics were contacted more frequently than those carrying rods marked by heterospecific females. Scent marks from conspecific females were also sniffed more frequently. A third experiment compared the level of responses to rods carrying material collected from the scent glands of female individuals to which the subjects had been habituated with their responses to rods carrying scent from females to which they had not been habituated. Contacting and sniffing responses to the scents of novel females were higher than those to the scents of females to which the subjects had been habituated.     

Olfactory Responses of Neotropical Short-Tailed Singing Mice, <Emphasis Type="Italic">Scotinomys teguina</Emphasis>, to Odors of the Mid-Ventral Sebaceous Gland: Discrimination of Conspecifics,Gender, and Female Reproductive Condition

We examined the olfactory communication of the Neotropical short-tailed singing mouse, Scotinomys teguina, by investigating whether S. teguina responded to odors produced by the mid-ventral sebaceous gland of conspecifics. Females spent significantly more time investigating male odor than an odorless stimulus or a female odor. Males spent significantly more time investigating female odor than an odorless stimulus, but not that of a male odor. This latter result does not seem to be explained by differences in age or sexual experience of test subjects, but may be influenced by reproductive condition of the female odor donor. Male S. teguina spent significantly more time (1) investigating and (2) in total proximity to odors of estrous than non-estrous females. Males spent more time (1) investigating and (2) in total proximity to odors of males than non-estrous females. Finally, given the choice between odor of males vs proestrous females, males showed no preference. Thus, the mid-ventral gland in S. teguina seems to convey information about conspecifics, sex, and female reproductive condition. Male odor, compared with proestrous female odor, is equally interesting to males, suggesting that the gland also plays an important role in male–male communication. Sexual dimorphism in the size of the gland and in the amount of secretion produced by the gland may be related to either male–male competition or female choice.     

Recognition of Dominance Status By Chemoreception in the Red Swamp Crayfish, Procambarus clarkii

We tested the hypothesis that chemical signals play a role in the recognition of dominance status in the red swamp crayfish, Procambarus clarkii. Dominance was judged on the outcome of dyadic interactions in all male or female groups of three individuals. This resulted in a dominant, intermediate, and subordinate individual within each population. A choice paradigm in a flow-through Y maze was used to judge whether crayfish were able to recognize dominance through chemical cues alone. Both individuals that interacted with the animal producing the odor and naive individuals were tested. Irrespective of sex and previous experience, individuals increased their rates of locomotion in the presence of conspecific odor. Naive males investigated the dominant arm first, spent more time at the dominant nozzle, and responded more aggressively (as measured by meral spread) to dominant male odor and subordinate female odor. Intermediate males spent more time at the dominant male nozzle and responded more aggressively to dominant male odor. Naive females spent more time at the dominant nozzle. These results show that males recognize dominant animals. Since both naive and experienced males respond to water from dominant animals, we concluded that this is recognition of dominance and not just individual recognition. This signal may be important for the formation or reinforcement of dominance relationships. Based on the change in behavior between odors, we suggest that crayfish can use chemical cues to recognize the dominance status of conspecifics.     

Urinary signaling pheromone and specific behavioral response in tree shrews (Tupaia belangeri)

As soon as they encounter the scent marks made by the urine of fertile male conspecifics, tree shrews of both sexes cover the scent marks with their own sternal-gland secretion by chinning. The urine of other species, female conspecifics, or castrated male conspecifics does not elicit this behavior; that is, overmarking with sternal-gland secretion is a reaction to a specific chemical signal. The intensity of the overmarking reaction, i.e., the chinning score in response to the urine sample, decreases as the urine is diluted. The urine of different male conspecifics elicits overmarking of varying intensity; an animal's own urine is in general not overmarked. This behavioral reaction meets all the criteria for a bioassay of potential use in isolating the tree-shrew pheromone from urine.     

Scent marks as social signals inGalago crassicaudatus II. Discrimination between individuals by scent

Thick-tailed galagos,Galago crassicaudatus argentatus, which had been habituated to the scent marks of a conspecific through repeated presentations, increased their sniffing when presented with scent marks from a second conspecific of the same sex. Thus, they discriminated between the scents of individual conspecifics. This result was obtained using naturally scent-marked perches and perches carrying only chest gland scent—the latter even two weeks after the marking. When urine, the prevalent scent signal among most other galago species, was tested, it elicited little interest and discriminations were not clearly made. It is suggested that the use of more specialized scents is related to the social characteristics ofGalago crassicaudatus.     

Chemically Mediated Mate Location and Evaluation in the Lobster, Homarus americanus

Many aspects of lobster (Homarus americanus) social behavior have been shown to be regulated by chemical signals. Female lobsters can determine the sex of sheltered conspecifics from a distance, and will approach and attempt to enter male-occupied shelters. We investigated (1) if female attraction to male-occupied shelters and entering decisions are based upon chemical signals, particularly male urine signals, and (2) if male dominance status and female molt stage influence these female behaviors. Sheltered dominant males were fitted with urine catheters, which collect urine and prevent its release into the environment. We found that males increased urine output in response to female entering attempts. Blocking male urine release significantly reduced the incidence of female shelter approach and the time spent attempting to enter. Artificial release of male urine in the presence of a catheterized male restored female approach but not time spent attempting to enter, while artificial urine release alone did not induce either behavior. Females approached any single male, but seemed to prefer to enter shelters occupied by more dominant males. Premolt and intermolt females differed in their behaviors: premolts showed more interest in approaching dominant male shelters without much effort to enter, while intermolts appeared less likely to approach but more motivated to attempt to enter shelters. This study suggests that both intermolt and premolt female lobsters use one or more chemical signals from sheltered males for location and evaluation of potential mates. Male urine appears to carry compound(s) important for both location and entering decisions, although nonurine chemical or other sensory signals are also implicated during close-up mate evaluation.     

Scent marks as social signals inGalago crassicaudatus I. Sex and reproductive status as factors in signals and responses

In simultaneous presentations of the scent marks of male and female conspecifics, thick-tailed galagos (Galago crassicaudatus) of both sexes responded most strongly to the female scent. Males differentiated between the scents most clearly in their sniffing, females in their scent marking in response. The scent of estrus females elicited increased licking of the scent by both sexes. This greater response to female scent and the contrasts in response patterns between the sexes are discussed and related to galago social and spatial relationships in the wild.     

Partial chemical characterization of urinary signaling pheromone in tree shrews (Tupaia belangen)

Tree shrews of both sexes exhibit marking behavior (chinning) in response to scent marks made by the urine of fertile male conspecifics. To isolate the effective odor components, the urine was fractionated by liquid-liquid extraction and TLC, and the fractions were tested by bioassay. The results show that chinning is elicited by several lipophilic urine fractions, which are more effective in combination than alone. To characterize the complex scent signal, the lipophilic extracts from urine of the two sexes were analyzed by GC-MS and compared. The GC profile of the males is distinguished by pyrazine compounds not detected in the profile of the females. The profiles of the sexes also differ with regard to several volatile monocarboxylic acids, which are present at higher concentrations in male than in female urine. More than 30 urine components have been identified. Synthetic equivalents of these urine components were bioassayed for effectiveness in eliciting chinning and compared with one another as well as with scent substances not normally present in tree-shrew urine. Strong chinning responses were elicited by (1) certain pyrazine compounds and (2) some monocarboxylic acids, when presented at the high concentrations specific to male urine. Marking behavior is usually not elicited by scent substances not contained in tree-shrew urine or by urine components common to mammals in general. The data so far available indicate that the male-specific scent signal of tree shrews is based less on a single unique component than on the qualitative and quantitative characteristics of a multicomponent mixture. In the bioassay, tree shrews of both sexes respond equally to the male specific substances by chinning. As the scent signal represented by these substances has a different meaning to males and females (rival or potential mate, respectively), chinning probably serves several different functions.     

Potential use of chemical cues for colony-mate recognition in the big brown bat,Eptesicus fuscus

Bats should benefit from recognition of their roost-mates when colonies form stable social units that persist over time. We used Y-maze experiments and gas chromatography–olfactometry (GC-O) to evaluate whether female big brown bats Eptesicus fuscus (Chiroptera: Vespertilionidae) use chemical cues to distinguish among conspecifics. In dual-choice Y-maze experiments, females chose the scent of another female from their own roost over a conspecific female from a different roost in a majority of trials. Analysis of total body odors using GC-O suggests that individuals from a given colony may share a more common odor signature with roost-mates than with non-roost-mate conspecifics. Using four principle components derived from 15 odor variables, discriminant function analysis correctly assigned most individuals to the correct colony.     

Patterns of deposition of urine containing chemosignals that affect puberty and reproduction by wild stock male and female house mice (Mus domesticus)

House mice release chemosignals in their urine that influence the timing of puberty and reproductive condition in conspecific females. These experiments tested the possibility that mice depositing urine containing chemosignals that affect puberty and reproduction do so differentially with respect to urine cues from conspecifics. Mice were tested in cages containing samples of urine or water on cotton in wire mesh capsules protruding from the cage floor. Their urine deposition patterns were recorded on squares of filter paper positioned below the cages. Males deposited more urine than females housed in groups, estrous females, diestrous females, or prepubertal females. All groups of mice deposited urine in a nonrandom fashion with regard to urine cues from conspecifics. Male mice deposited more urine near locations previously soiled by females than near water or other males. Grouped females deposited more urine near male urine cues and avoided depositing urine near urine from other group-housed females. Both estrous and diestrous female mice deposited more urine near males than near other urine cues or water, possibly to attract mates. Prepubertal females avoided depositing their urine near male urine and urinated more near urine from grouped females than near other urine types or water. Young females may be avoiding possible male mates until they have attained puberty. This avoidance behavior may enhance the long-term reproductive success of the females that otherwise might mature and mate at too young an age.     

Scented Males and Choosy Females: Does Male Odor Influence Female Mate Choice in the Mediterranean Fruit Fly?

The Mediterranean fruit fly, Ceratitis capitata (Wiedemann), displays a lek mating system characterized by a high level of female discrimination among potential mates. The basis of female choice is not understood, but recent studies indicate that male exposure to the aroma of certain plant structures or essential oils may increase mating success. In particular, exposure to the aroma of ginger root oil (GRO) enhances male mating frequency, and several sterile-male release programs against C. capitata have incorporated ‘aromatherapy’ (large-scale exposure of pre-release insects to GRO) to increase the effectiveness of control efforts. We investigated the mechanism underlying female preference for GRO-exposed males. Two sets of experiments were conducted. In the first, we monitored female attraction to (1) freshly killed flies, or (2) paper discs that contained hexane extracts from varying treatments. In these tests, females were sighted more often (1) near GRO-exposed than non-exposed males (even when the males were visually concealed) and (2) near extracts from GRO-exposed than non-exposed males. These findings suggest a ‘perfume effect’, whereby female mate choice is mediated by olfactory differences. In the second set, we compared (1) mate choice between intact females and females from which both antennae had been surgically removed, and (2) mating success between intact males and males from which both antennae had been surgically removed before GRO exposure. Intact females preferred GRO-exposed males, whereas females lacking both antennae rarely mated and showed no preference between GRO-exposed and non-exposed males. In the opposite treatment (intact females but surgically altered males), GRO-exposed males lacking both antennae mated as frequently as GRO-exposed intact males. These data suggest that female choice was dependent on olfactory perception of male odor but that male mating success did not depend on olfactory perception of GRO aroma, suggesting, in turn, that GRO conferred a mating advantage through an external phenomenon (possibly alteration of cuticular scent) rather than through internal processing (pheromone synthesis).     

Use of Chemical Communication by the Subterranean Rodent Ctenomys talarum (Tuco-Tuco) During the Breeding Season

Solitary subterranean rodents with a low frequency of direct contact between conspecifics are expected to use chemical communication to coordinate social and reproductive behavior. We examined whether reproductive tuco-tucos (Ctenomys talarum) were able to discriminate the reproductive condition, sex, and source population of conspecifics by means of chemical cues contained in urine, feces, soiled shavings, or anogenital secretions. During preference tests in which animals had direct contact with these chemical cues, tuco-tucos were able to determine the reproductive condition of opposite sex conspecifics independent of the source of odor. When only olfactory cues were available, both sexes discriminated reproductive condition of opposite sex individuals using urine. Females were also able to discriminate the reproductive condition of males using soiled shavings. Females spent more time investigating male odors than female odors; except in the case of feces, breeding males spent similar amounts of time investigating male and female odors. No preferences were detected for opposite sex urine from members of an animal's own versus another population. The role of chemical cues in territory defense and breeding performance by this highly territorial subterranean rodent is discussed.     

Conspecific scent trailing by garter snakes (Thamnophis sirtalis) during autumn Further evidence for use of pheromones in den location

Adult garter snakes (Thamnophis sirtalis), collected in October near a traditional, communal hibernaculum in central Wisconsin, were acclimated to autumnal conditions and subjected to laboratory tests to determine whether they could follow scent trails of a conspecific. Graded responses were obtained, but 75% of the sample showed at least some inclination to follow scent trails. The results suggest that pheromone cues may be used by male and female garter snakes to locate traditional dens during autumnal migrations. Evidence from this and other studies suggests that pheromones are probably used in conjunction with other homing mechanisms and that the role of pheromones in den location may be more important in younger snakes and in populations inhabiting northern latitudes.     

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.     

Volatile components of scent material from cotton-top tamarin (Saguinus o. oedipus)

The first chemical analysis of the volatile components of scent material from the cotton-top tamarin,Saguinus o. oedipus is presented. In contrast to our previous findings in saddle-back tamarins (Saguinus fuscicollis), the chemical composition of the scent marks is quite variable. Analysis of secretion expressed manually from different areas of the scent pads of sedated animals shows individual and regional variability. Only three components (squalene, cholesterol, andp-methoxybenzaldehyde) were found in all scent mark samples analyzed. However, suprapubic secretions obtained from one sedated female contained, in addition, 12 butyrate esters as well as five acetate esters which were homologous to the butyrates. The butyrate esters have been previously identified in scent material fromSaguinus fuscicollis. The variability of the composition of the scent material is discussed in relation to the scent-marking behavior of the cotton-top tamarins. Behavioral studies tested the ability of the cotton-top tamarins to discriminate between scent marks from conspecifics and scent marks from saddle-back tamarins. The group of 12 subjects discriminated between the scent marks from both species in choice tests during which they could freely contact the scent samples. However, when scent marks were presented under a screen, so that the subjects could smell but not contact the stimuli, no discrimination was shown. Four individuals displayed more interest in the screened scent stimuli than all other subjects. When these animals were tested in a second experiment for their ability to discriminate between material from the two species on the basis of only volatile cues, it became evident that they were able to do so. These results indicate that volatile cues alone enable the tamarins to recognize scent marks from conspecifics, but that additional cues perceived during contact with the scent are important for its full attractiveness and/or informational content.     

Volatile compounds from the predatory insectPodisus maculiventris (Hemiptera: Pentatomidae)

In the predatory spined soldier bug,Podisus maculiventris, the dorsal abdominal glands are much smaller in adult females than males. Females produce a mixture of (E)-2-hexenal, (E)-2-octenal, (E)-2-hexenoic acid, benzaldehyde, and nonanal in these glands. The female dorsal abdominal gland secretion may be a close-range pheromone since the dorsal abdominal gland secretion from males has been shown to be a long-range aggregation pheromone in this species. The metathoracic scent gland secretions of male and female spined soldier bugs are apparently identical, and similar to that of other pentatomids, except for the presence of the monoterpene alcohol, linalool.Mention of a commercial product does not constitute an endorsement of this product by the USDA.     

Intersexual Differences in Chemical Composition of Precloacal Gland Secretions of the Amphisbaenian <Emphasis Type="Italic">Blanus cinereus</Emphasis>

Pheromonal communication seems to be especially important in amphisbaenians, a group of reptiles morphologically and functionally adapted to fossorial life. Both male and female amphisbaenians (Blanus cinereus) produce copious amounts of a secretion from the precloacal glands. Analysis by gas chromatography–mass spectrometry revealed that these secretions contain 29 major lipophilic compounds, including several steroids (mainly cholesterol and cholesteryl methyl ether), n-C₉ to n-C₁₈ carboxylic acids (more diverse in males), and methyl and long-chain (waxy)-type esters of carboxylic acids, along with squalene (mainly in males) and tocopherol (only in females). There were clear intersexual differences in the presence/absence of some compounds, such as some fatty acids, steroids, and tocopherol, and in the relative proportions of some shared compounds, such as squalene. These differences may explain how the sexes of this amphisbaenian discriminate one another based on scent alone. The abundance of steroids and waxy esters in secretions of both male and female amphisbaenians may be useful to scent mark underground tunnels to facilitate orientation by these organisms.     

Male scent-producing structures inColias butterflies

During successful courtship in sulfur butterflies, virgin females respond to males by assuming a stationary posture and extending the abdomen ventrally from between the hindwings, thereby permitting copulation. In the clouded sulfur,Colias philodice Godard, this response was used in a laboratory bioassay to confirm the existence of a male chemical signal demonstrated by a previous worker and to document the signal's behavioral function and source. The male scent is shown to be required, in part, to reliably elicit abdominal extension and to be emitted from a patch of cells and scales associated with the dorsal surface of the male hind wing near the wing base. Experiments also show that evaporation of the signal is reduced when the source is covered by the forewing as it is at rest and in flight. These data, coupled with other information on male chemical signals in sulfur butterflies and other Lepidoptera, suggest that in sulfurs the morphology and chemistry of the scent glands along with the behavior of the male are structured in a way that minimizes the evaporative loss of scent from the wings when the male is not courting females.     

Male bugs modulate pheromone emission in response to vibratory signals from conspecifics

Male Nezara viridula emit a volatile sex pheromone that acts as a long-range attractant to females. Both sexes communicate through vibrations once they are on the same plant. Males respond to the female calling song by emitting a male courtship song, and they orient to the female calling song on a plant. Simultaneity between vibratory and chemical communication during the last steps of mate finding suggests that pheromone emission might be modulated by signals from conspecifics. Male volatile emissions were collected with fibers for solid-phase microextraction, while male bugs were stimulated with natural and artificial signals. Percentages of males releasing pheromone and collected amounts of pheromone increased when males were stimulated with female calling song. Pheromone emission was stable in males stimulated with male rivalry songs, and it decreased in males stimulated with a 100-Hz artificial signal. The ability of male bugs to modulate their pheromone emission may reduce metabolic costs, reduce parasitism, and offer a better synchronization of sexual activity.     

Increased scent marking in male Mongolian gerbils by urinary polypeptides of female conspecifics

Male Mongolian gerbils scent mark during a standardized openfield test on an individually different but more-or-less constant level in the absence of females. The presence of females in the same room but without direct contact to the males increases the marking frequencies of the males to an individually different higher level. A similar increase in marking frequency is obtained by application of female urine directly to the nostrils of the males, but not by male gerbil urine or female laboratory mouse urine. This indicates that substances present in female gerbil urine are responsible for the increase in male marking behavior. Using this bioassay, various fractions of female urine were tested for their influence on marking behavior. Our results to date show that the component increasing male scent-marking behavior is hydrophilic and nonvolatile and is contained in the nondialyzable, high-molecular-weight polypeptide fraction of female gerbil urine.