Selection of scent-marking sites by klipspringers (Oreotragus oreotragus)

Klipspringers deposit scent marks in the form of secretion from the preorbital glands on branches of low trees and shrubs within their territories. Local environmental factors which affect choice of scent-marking sites in a population of wild klipspringers in Zimbabwe are described. Scent marks were typically placed within a narrow vertical range at the end of upward-pointing twigs. Twigs were more likely to be selected if they were located above bare rock, particularly if the ground was raised above the immediately surrounding area or was close to a significant break in slope. A substantial proportion of sites were placed on dead wood. Marking sites may remain in use for a number of years, with turnover rates calculated at between 1 and 2% per annum. The species of tree on which marks were placed were recorded and compared with overall abundance in the habitat. Palatable tree species were found to be selected preferentially when compared to unpalatable ones. Several explanations for this are compared. It is concluded that, simply through preferential selection of certain local characteristics, klipspringers may enhance the visibility and predictability of their scent marks.     

Social Organization and Ecology of the Klipspringer (Oreotragus oreotragus) in Ethiopia

Der Klippspringer (Oreotragus oreotragus) wurde in drei Gebieten in Athiopien beobachtet. Meist leben ein ♂ und ein bis zwei ♀♀ zusammen. Die Gruppe hat ein kleines Wohngebiet (etwa 8 ha), das sie gegen andere Gruppen verteidigt. Klippspringer fressen meist Gras und Blätter von Büschen. Klippspringer haben viele Feinde. Das ♂ steht meist Wache, während das ♀ gesichert frißt. Das ♀ führt Ortswechsel an. Die Jungen werden nach der Regenzeit im November geboren. Im folgenden Jahr verlassen die männlichen Jungen ihre elterliche Gruppe; die weiblichen Jungen bleiben länger darin, und einige paaren sich mit ihrem Vater. Stabile Paarbindungen werden als Anpassung an offene Landschaft gedeutet; das ♂ kann ♀♀ und Junge beschiitzen.     

Behavioural Responses to Scent Marks of Increasing Age in Klipspringer Oreotragus oreotragus

Klipspringers Oreotragus oreotragus are small antelopes which use scent marks to demarcate and defend their territory. The strength of behavioural response to the introduction of unfamiliar scent was measured in order to estimate the effective lifespan of scent secretion. The total number of scent marks deposited during the experiment was higher than control levels when fresh scent was introduced, but not when older scent was introduced. However, the proportion of marks deposited on introduced twigs remained higher than control levels until scent was aged at least 7d. The probability of licking twigs immediately before scent-marking was much higher than control levels when scent aged up to 3d was introduced, but not when scent was aged 7d or more. These results indicate that preorbital gland scent retains at least some degree of efficacy for up to 7d. The decreasing level of response to scent aged 0-7d is most likely to be due to decay of the scent signal. However, in contrast to the total number of marks deposited and the probability of licking before scent-marking, which both consistently decreased in response to older scent, the proportion of marks deposited on scent aged 3d was higher than over fresh scent. It is suggested that, in view of the role that scent-marking plays in territory defence, this difference in the timing of peak response between the three variables may represent an adaptive response to the perceived threat of encounter with an adversary, which is mediated by the age of the scent marks.     

Spatial and temporal scaling in habitat utilization by klipspringers (Oreotragus oreotragus) determined using giving‐up densities

An animal's pattern of habitat use can reveal how different parts of its environment vary in quality based on the costs (such as predation risk) and benefits (such as food intake) of using each habitat. We studied klipspringer habitat use in Augrabies Falls National Park, South Africa using giving‐up densities (GUDs; the amount of food remaining in a resource patch following exploitation) in experimental food patches. We tested hypotheses related to how salient habitat variables might influence klipspringers' perceptions of foraging costs. At small spatial scales (3–4 m), klipspringer GUDs did not vary with cover and open microhabitats, or with the four cardinal aspects (shading) around shrubs. Adding water adjacent to food patches did not influence GUDs, showing that water is not a limiting complementary resource to food. Generally, klipspringers do not appear to be physiologically constrained. There was no difference in GUDs between four daily time periods, or between summer and winter; however, a significant interaction effect of time‐of‐day with season resulted from GUDs during the midday time period in winter being significantly higher (perceived value lower) than during the same time period in summer. At moderate spatial scales (10–60 m), klipspringer GUDs increased with distance from rocks because of increased predation risk. Based on GUDs collected at the largest scale (two 4.41‐ha grids), klipspringers preferred foraging at greater distances from drainage lines and on pebble and cobble substrates. Overall, this study has shown the efficacy of measuring GUDs to determine klipspringers' habitat utilization while foraging.     

Species and subspecies specificity in urine and scent marks of saddle-back tamarins (Saguinus fuscicollis)

Saguinus fuscicollisproduces scent marks which consist mainly of a mixture of urine and the secretions of circumgenital scent glands. The present study investigates the ability of saddle-back tamarins to discriminate between scent material from conspecifics and corresponding material from other species and to differentiate material from two subspecies of Saguinus fuscicollis.When choices between urine samples from conspecifics and from guinea pigs and choices between urine samples from conspecifics and from common marmosets were offered, the tamarins investigated samples from conspecifics more frequently. Similar responses were obtained when choices between scent marks from saddle-back tamarins and from common marmosets and between scent marks from saddle-back and red-chested moustached tamarins were offered. The tamarins also discriminated between scent marks and between extracts of scent marks from Saguinus f. fuscicollisand Saguinus f. illigeri.In these choice tests, subjects of both subspecies tended preferentially to investigate material from Saguinus f. fuscicollis.The results of these studies show that urine and scent marks contain chemical cues on which recognition of conspecifics can be based. Moreover, the scent marks of closely related subspecies also offer cues which could enable the tamarins to discriminate between them.     

Sex and age-specific differences in ultraviolet reflectance of scent marks of bank voles (Clethrionomys glareolus)

Scent markings of voles are visible via their ultraviolet reflection. Kestrels, and possibly other diurnal raptors, may use this property when hunting. We performed a laboratory study on bank voles to determine whether UV-reflectance of scent marks differs in relation to sex, age and social status. When reflectance spectra of scent marks were measured with a spectroradiometer, we found UV reflectance to be strongest in mature males. There were no differences between mature females and immature juveniles, nor between sexes in juveniles or mature and immature individuals in females. Moreover, we did not find any difference in UV reflectance between dominant and subordinate mature males. The results of this study support earlier findings that UV sensitive predators may use UV reflectance of scent marks as a prey cue. Consequently, studies on differing vulnerability of voles to avian predators should take into account not only their space use and behaviour but also the UV reflectance of their scent marks. Accepted: 21 August 1999     

Proteins: biologically relevant components of the scent marks of a primate (Saguinus fuscicollis)

We report here the first documentation that proteins contributeto the scent communication system in a primate. Data presentedhere indicate that high molecular weight compounds which arenot airborne are required for perception of the full attractivenessof the scent. Scent marks from the saddle-back tamarin, a SouthAmerican primate, contain a number of water-soluble proteins.Gel electrophoresis documents that the major protein (66 kD)is found in urine, and a second protein (18 kD) comes from thegland secretions. The 18 kD protein produces two distinct bands(14 kD and 4 kD) under reducing conditions. Concurrent behavioralstudies demonstrate that removal of proteins by enzymatic degradationdoes not alter the ability of the tamarins to discriminate betweenscents from different donor types. However, subtle changes insensory quality occur when the proteins have been degraded.In particular, tamarins do discriminate material in which theproteins have been digested from material in which the proteinsare intact. This observation suggests that proteins are a componentof what constitutes the scent image. In addition, they may serveas carriers and/or reservoirs for more volatile ligands thatencode some of the communicatory messages contained in the material.     

The ownership signature in mouse scent marks is involatile

Male house mice advertise their territory ownership through urinary scent marks and use individual-specific patterns of major urinary proteins (MUPs) to discriminate between their own scent and that of other males. It is not clear whether recognition occurs through discrimination of the non-volatile proteins or protein-ligand complexes (direct model), or by the detection of volatile ligands that are released from MUPs (indirect model). To examine the mechanism underlying individual scent mark signatures, we compared investigatory and countermarking responses of male laboratory mice presented with male scent marks from a strain with a different MUP pattern, when they could contact the scent or when contact was prevented by a porous nitrocellulose sheet to which proteins bind. Mice investigated scent marks from other males whether these were covered or not, and biochemical analysis confirmed that the porous cover did not prevent the release of volatiles from scent marks. Having gained information through investigation, mice increased their own scent marking only if they had direct contact with another male's urine, failing to do this when contact was prevented. Individual signatures in scent marks thus appear to be carried by non-volatile proteins or by non-volatile protein-ligand complexes, rather than by volatiles emanating from the scent.     

王朗自然保护区中大熊猫发情场的嗅味树和嗅味标记调查

2004年2月底至4月中旬,我们对四川省王朗国家自然保护区内大熊猫(Ailuropodamelanoleuca)发情场的嗅味标记和嗅味树进行了调查,以了解王朗大熊猫的发情时间和发情场位置,为确定保护区内的重点保护区域和时间段提供证据,同时增加对大熊猫生物学特性的了解。在海拔2500–3200m间,共对10条样带进行了36次调查,测量并记录了山脊上1801株乔木的种类、高度、胸径和嗅味标记状况。调查共发现24株被大熊猫标记的嗅味树,其中54%位于种群分布中心区域内两侧都有森林分布的山脊上,70.8%分布于海拔2700–3000m之间。嗅味树的平均胸径为37.2±17.8cm(N=24),其中92%为岷江冷杉(Abiesfargesiivar.faxoniana),96%树皮粗糙或比较粗糙。嗅味标记中,75%的嗅味标记为肛周腺分泌物,标记中心的平均高度为51.8±15.6cm(N=18),25%为尿液,尿液上沿的平均高度为64.5±14.5cm(N=6)。此外,从调查结果初步得知,春季王朗最重要的大熊猫发情场所是长白沟-解放沟等区域,这里是大熊猫种群的关键栖息地。目前保护区对这些关键栖息地加强了保护力度。     

Do scent marks increase predation risk of microtine rodents?

Reproductive activities, including signalling with scents, may increase the risk of predation. Mammalian predators, like small mustelids, find voles by using odour cues of scent marks. Scent marks are also visible in ultraviolet light, and at least some diurnal raptors are attracted to these markings in the field. We performed a field experiment to find out whether manipulation of scent markings affects the density, survival and mobility of free-living voles, and the activity of mammalian and avian predators. A total of 20 plots (each 1 ha) were randomly divided into two treatments: scent manipulation and control plots. Scent manipulation plots were treated with vole scent-liquid and control plots with water. From each plot 1–2 voles were radio-collared and tracked for three weeks. Predators hunted more often on scent manipulation plots than on control plots leading to lower survival time of voles. Although scent manipulation attracted more avian predators, small mustelids were the main predators of voles. The density or the mobility of voles did not differ significantly between manipulation and control plots. Our results suggest that odour of scent marks may be a larger risk to voles than UV visibility of scent marks.     

Recognition of scent marks in solitary bees to avoid previously visited flowers

Knowing how floral visitors forage efficiently among flowers is important to understanding plant-pollinator interactions. When bees search for rewarding flowers, they use several visual cues to detect the available floral resources. In addition to these cues, bees can recognize scent marks, which are olfactory cues left on flowers foraged by previous visitors. This behavior is well known in social bees, such as honeybees and bumblebees. Although solitary bees do not need to give information about which flowers were foraged to conspecifics, several pieces of evidence have indicated the use of scent marks. However, it is unknown whether the behavior is widely used in many different bee species. We investigated whether four different solitary bees, Colletes patellatus (Colletidae), Andrena prostomias (Andrenidae), Osmia orientalis (Megachilidae), and Tetralonia mitsukurii (Apidae), can recognize flowers that have been foraged previously by visitors within 3 min. All four bees showed rejection responses to flowers foraged by conspecifics. However, our results showed that responses to foraged flowers varied among bee species. The tendency of A. prostomias and T. mitsukurii to reject the foraged flowers was pronounced, while in C. patellatus and O. orientalis it was weak. In both A. prostomias and T. mitsukurii, the rejection rate of flowers foraged by conspecifics decreased as the time lag after the last visit increased. Both bees visited the flowers from which pollen or nectar had been artificially removed. We suggest that A. prostomias and T. mitsukurii would recognize scent marks left by previous visitors, while the other two bees would not recognize them so strongly. It is likely that the decision to use scent marks is dependent either on the richness of resources or on the complexity of floral structure.     

Red foxes (Vulpes vulpes) use rabbit (Oryctolagus cuniculus) scent marks as territorial marking sites

One of the functions of chemical communication is territorial signalling. To achieve this, animals should mark their territories in a manner that increases the detectability of the marks, thereby maximising the probability that other animals detect the scent marks. In this study, we focused on the scent-marking behaviour of red foxes in relation to the abundance of their main prey, the European rabbit, in a suburban forest in Madrid, Spain. Our results reveal that foxes scent-marked more and increased the detectability of their marks in areas of higher rabbit density. It would appear that foxes defend food resources from competitors by increasing the number and the detectability of their scent marks.     

Female marmoset monkeys (Callithrix jacchus) can be identified from the chemical composition of their scent marks.

The present study analyzed 42 organic solvent extracts of scent mark pools from five dominant female common marmosets by gas chromatography (GC) and combined GC and mass spectrometry. We determined whether there were qualitative or quantitative differences between the chemical composition of scent marks from individual females. Gas chromatography and mass spectral analysis detected the same 162 chemicals in 86% (36/42) of scent mark pools from five dominant females. This near identical chemical composition of scent marks suggested there were few, if any, qualitative differences between the chemical composition of scent marks from individual females. Instead, quantitative differences in scent may provide the key factor distinguishing individual females. Using the relative concentration of highly volatile chemicals detected by GC in scent marks, linear discriminant analysis classified scent mark pools to their correct donor approximately 91% of the time. Such highly reliable statistical matching of scent to donor suggested that each individual female common marmoset has a unique ratio of highly volatile chemicals in their scent marks which may permit individual identification of females from odors in their scent alone.     

Discrimination by male ring-tailed Lemurs (Lemur catta) between the scent marks of male and those of female conspecifics

Olfaction plays an important role in the social communication of all prosimians. (The experiment reported in this paper forms part of an intensive chemobehavioral study of olfaction in Lemur catta (ring-tailed lemur) being carried out in this laboratory.) Five male Lemur cattawere tested on their behavioral responses to paired scent stimuli. Responses measured were (1) total investigation time, (2) arm-marking, (3) ABO/BO rubbing, and (4) flehmen. Males showed a strong discrimination between the scent stimuli,giving higher levels of response to female scent on measures 1, 3, and 4. This response suggests an olfactory-related preference by males for female scent under controlled conditions. This preference may be a consequence of the females’ dominance over males and the brevity of estrus in L. catta,both of which would favor such choice behavior.     

Floral scent and intrafloral scent differentiation inMoneses andPyrola (Pyrolaceae)

Floral scent was collected by headspace methods from intact flowers, petals, and stamens of four species ofPyrolaceae. The scent samples were analyzed by coupled gas chromatography-mass spectrometry (GC-MS). The floral scent inPyrola spp. is differentiated into a characteristic petal scent—phenyl propanoids and a characteristic stamen scent—methoxy benzenes. InMoneses the scent is characterized by isoprenoids and benzenoids, with a larger proportion of benzenoids in the stamens compared to the petals. Specific anther scents may promote foraging efficiency in buzz-pollinated species and enhance flower fidelity. Variation in floral scent composition is consistent with the taxonomic relationships among the genera and species examined.     

Chemical composition of scent marks in the ringtailed lemur (Lemur catta): glandular differences, seasonal variation, and individual signatures

The apocrine and sebaceous scent glands of ringtailed lemurs (Lemur catta) appear to serve different social functions. In behavioral experiments, lemurs modulate their responses to scent marks based on the type of odorant, their own physiological state, the signaler's physiological state, and prior social experience. To examine variation in odorant chemistry relative to olfactory behavior, we used gas chromatography and mass spectrometry to analyze over 86 samples of glandular secretion collected over 2 years from 15 adult lemurs. Labial and scrotal secretions contained organic acids and esters, whereas male brachial secretions were composed almost entirely of squalene and cholesterol derivatives. Principal component and linear discriminant analyses revealed glandular, individual-specific, and seasonal variation in chemical profiles but no relationship to the signaler's social status. The chemical composition of the various secretions provides further clues about the function of the different glands: the higher molecular weight compounds in genital and brachial secretions may increase signal longevity and provide lasting information to conspecifics, consistent with a role in advertising resource ownership or reproductive state. Conversely, the lower molecular weight compounds of antebrachial secretions produce ephemeral signals used primarily in social dominance displays and require integration of multiple sensory modalities for effective signal transmission.     

Sniffing out patterns of sexual dimorphism in floral scent

1.   A major transition in flowering plants has been the evolution of separate sexes (dioecy) from combined sexes (hermaphroditism). This transition is often, but not always, accompanied by the evolution of sexual dimorphism in attractive traits, and floral scent is no exception.
2.   In this review I aim to improve our understanding of variation in sexual dimorphism in floral scent characteristics by first explicating the relevant hypotheses, and then deriving explicit predictions for the pattern of floral scent from each.
3.   Next, I synthesize and qualitatively review published data on floral volatile emission rate and composition in 33 gender dimorphic species to identify emerging patterns, and evaluate these in light of predictions derived from the hypotheses.
4.   Although conclusions must be viewed as preliminary, a handful of strong patterns were revealed: (1) in the majority of the species studied males emit more volatiles per flower than females, (2) in over half of the species studied the sexes differed in at least one aspect of scent composition, (3) sexual dimorphism in scent composition was less common in species with rewardless females and/or brood site pollination than those that offered nectar and/or pollen, (4) a one-to-one correspondence between sex differences in overall scent composition and male organ-specific scent production is largely not found.
5.   This review has highlighted gaps in our understanding of the genesis of patterns of sexual dimorphism in floral scent, and makes clear that to move the field forward we need to shift our focus from pattern to process, and this will be best achieved by simultaneously testing alternative hypotheses at the same level of analysis.