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MAIK “Nauka

/Interperiodica”0584

Biology Bulletin, Vol. 30, No. 6, 2003, pp. 584–590. Translated from Izvestiya Akademii Nauk, Seriya Biologicheskaya, No. 6, 2003, pp. 698–705.Original Russian Text Copyright © 2003 by V. Rozhnov, Yu. Rozhnov.

The common palm civet (

Paradoxurus hermaphro-ditus

Pallas, 1777) is a convenient model species forstudying mediated communication by scent marks intropical small carnivorous mammals. As many of theseanimals, this species has a specific skin gland, theperineal gland. The secretion of this well-studied gland(Pocock, 1915; Sokolov and Neklyudova, 1988) can beused for mediated communication by scent marks.These animals also leave urine and feces as scent marks(Rozhnov

et al.

, 1992). However, the role of each ofexcretion in mediated communication by scent marksof the common palm civet remains unclear. Previously(Rozhnov and Rozhnov, 1998) we presented the datademonstrating different olfactory responses of thesetropical carnivorous mammals to their urine and feces.

The goals of this work were as follows: (1) to studythe marking behavior and activity in leaving scentmarks of different origin (perineal gland secretion,urine, and feces) in the common palm civet, (2) to com-pare the olfactory responses of these animals to thereleased excretions that can be used as scent marks inscent-mediated communication, and (3) to estimatetheir ability to distinguish the animals of their own andother species, males and females, as well as familiarand unfamiliar individuals.

MATERIALS AND METHODS

The experiments were carried out at the Tcher-nogolovka Biological Station (Severtsov Institute ofEcology and Evolution, Russian Academy of Sciences)

in June–September 1996 and 1997. We used four adultmales and two adult females in 1996 and six adultmales and two adult females in 1997. They were theoffsprings of animals brought from Vietnam (one adultfemale was provided by the Leningrad Zoo, St. Peters-burg). The animals were kept individually, in hetero-sexual pairs, or unisexual groups of 2–3 specimens inopen-air wire cages (

1.8

×

1.8

×

2.2, 1.8

×

3.6

×

2.2

, or

3

×

6

×

4

m) with a complex interior that allowed theanimals to use the entire three-dimensional space of acage.

The frequency of scent marking was determined byvisual observation of the animals for 1–4 h a day(depending on whether conditions) during their loco-motor activity. Observations continued for a total of114 h.

Information contents of the excretions were evalu-ated by the method of two-choice situation previouslyused in the experiments with other small carnivorousmammals (Rozhnov, 1979; Sokolov and Rozhnov,1983, 1989; Rozhnov and Rogoschik, 1994). A pair ofodor sources differing in a single parameter (species,sex, or familiarity) was simultaneously presented to arecipient. Each of the studied odors was applied to apredeodorized tinplate. The plates were placed on acage wall at the level of animal nose at a distance of 5–10 cm. The duration of animal sniffing of each odorsource was recorded using two Slava start/stopwatches. The experiment was carried out until theresponse to odor faded (an animal stopped sniffing theplates with odors). Only one pair of odors was pre-

Roles of Different Types of Excretions in Mediated Communication by Scent Marks of the Common Palm Civet,

Paradoxurus hermaphroditus

Pallas, 1777 (Mammalia, Carnivora)

V. V. Rozhnov and Yu. V. Rozhnov

Severtsov Institute of Ecology and Evolution, Russian Academy of Sciences, Leninskii pr. 33, Moscow, 119071 Russia

e-mail: rozhnov@sevin.ru

Received February 24, 2000

Abstract

—We studied scent-marking behavior, olfactory response to various excretions (secretion of theperineal gland, urine, and feces), as well as the information content of these excretions in the common palmcivet (

Paradoxurus hermaphroditus

Pallas, 1777). Six forms of scent-marking behavior have been described.The activity and proportion of forms of such behavior differ in males and females. Dragging the perineal glandand leaving the secretion on the substrate was most commonly observed in the animals of both sexes. The olfac-tory response of the palm civet varied in duration and depended both on animal sex and excretion type. Thepalm civet could distinguish animal species, sex, and familiar/unfamiliar individuals by the odor of the perinealgland secretion. Differences in the olfactory response of the palm civet to feces was observed only if theybelonged to different species. The scent marks of the perineal gland were most significant in mediated commu-nication by scent marks of the palm civet; they completely replicated the information of the urine marks.

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sented to an animal per day, and the position of theplates on a cage wall was chosen at random. The exper-iment was not taken into account if a recipient sniffedonly one odor source or if the difference in the durationof sniffing was less than 0.2 s (this value corresponds toa possible observation error). The fact that the pre-sented odor sources conveyed valid information wasconfirmed by (i) the asymmetric olfactory response ofthe animals to the presented two odor sources, whichwas manifested as directed olfactory response to one ofthe two odors and (ii) different durations of sniffing. Atotal of 396 presentations have been made. The experi-ments on the ability of the palm civet to distinguishtheir own species from another one were carried outusing the perineal gland secretion of a male forest genet(

Genetta pardina

) and the urine and feces of a maledomestic cat (

Felis domestica

). The feces and urinewere collected in a

50

×

25

×

25

-cm welded mesh cagewith a mesh size of

24

×

24

mm mesh size, while theperineal gland secretion was collected in a speciallydesigned cage of the same size with a locking ceiling.The urine and feces were collected several hours priorto the experiment, and the perineal gland secretion,immediately before the experiment.

Experimental data were processed statistically usingthe nonparametric sign test and standard statistical tests(Zaitsev, 1973; Lakin, 1990).

RESULTS AND DISCUSSION

Scent-Marking Behavior of the Palm Civet

Below, we describe the following forms of scent-marking behavior recorded during our observations:dragging the perineal gland, rubbing the ear–neck area,heel rubbing, anal drag, urination, and defecation.

1. Dragging the perineal gland.

This is a conspic-uous form of scent marking that can be observed whenthe palm civets are on the ground, trees, shelters hang-ing on the cage walls, etc. An animal squats, bending allfour legs and pressing the posterior abdomen (theregion of the perineal gland) against the substrate, andmakes a sliding movement forward. While on theground, the palm civet prefers to drag over prominentobjects (roots, large branches lying on the ground, etc.);however, the position of the gland allows an animal toleave the secretion on a flat surface as well. Usually ani-mals use several permanent places of marking by theperineal gland secretion, so that such marked objectsare clearly visible and have a specific smell.

2. Rubbing the ear–neck area.

The palm civetdemonstrates this way of scent marking both on theground and objects rising above its surface. An animalpresses the upper cheek–ear region against the substrateand slides forward, so that the neck and sometimes evenbody side are involved in rubbing. While moving alongtree branches in the cage, the palm civet rubs againstthese branches, cage walls, shelters hanging on them,etc. While on the ground, an animal prefers tree trunks,

cage walls, and other prominent objects (e.g., stumps)for rubbing.

3. Heel rubbing

is usually observed when the ani-mals are on the ground, but they also display this behav-ior on wide above-ground surfaces, such as the roofs ofshelters or thick logs and branches. An animal squats onits hind legs and “wipes” the soles on the substrate,moving them alternately forward. This is accompaniedby a typical shuffling sound.

4. Anal drag.

The animals usually display thisbehavior immediately after defecation, either on theground or on the substrates above the ground. An ani-mal arches its back, squats on its hind legs, “tucks in”its rump, and slides forward, dragging the anal regionover the substrate and sometimes “walking” on fore-feet.

5. Urination.

Palm civets usually urinate on theground. Performing this vital function, an animalslightly squats, spreads its hind legs, and holds the tailhorizontally or obliquely, at a sharp angle. Urination ina similar posture while moving forward, which resultsin a “urine path left on the ground, was described for thepalm civet; however, no such behavior has beenobserved during the study period.

6. Defecation

was observed on the ground, treebranches, and house roofs. Usually animals defecatedin the same special places, organizing “latrines” (indi-vidual or communal) with a large amount of feces,although single portions of feces could be elsewhere.During defecation, an animal arches its back andslightly squats; at the same time, it can move forward,shifting all four legs and holding its tail at an angle tothe horizontal.

The proportions between the above-mentionedforms of the palm civet behavior are given in Fig. 1.Dragging the perineal gland and rubbing the ear–neckarea are most common in both males and females. Inaddition, males often rub their heels, while such behav-ior is less frequent in females. Anal drag, urination, anddefecation have similar proportions in scent-markingbehavior of males and females, although anal drag isslightly more frequent in females.

123456

34 5 6

12

14%6% 2%1%

55%22%

1

23 4 5 6

24%2% 2%4% 2%

66%

Fig. 1.

Ratios of the forms of scent-marking behavior in male(left) and female (right) palm civets: (

1

) dragging the perinealgland, (

2

) rubbing the ear-neck area, (

3

) heel rubbing, (

4

) analdrag, (

5

) urination, and (

6

) defecation. The sum of all forms ofbehavior (

n

= 981 in males and

n

= 207 in females) is taken as100%.

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Table 1 presents the data on scent-marking activityin males and females. No significant quantitative differ-ences in the frequencies of different forms of scentmarking in males and females have been revealed.Note, however, that the confidence levels for heel rub-bing and urination frequency were close to the thresh-old. Sex-related differences in the proportions of theseforms of behavior in the palm civet are given in Fig. 2.

Olfactory Response of the Palm Civet to Excretions of Various Types

The olfactory response of the palm civet to the pre-sented odor sources is typical: an animal approaches anodor source and sniffs, placing its nose very close to it.Usually the animals sniffed a presented pair of odorsources once and then went away. The second sniffing(if at all) was considerably shorter than the first one.

The minimum and maximum recorded durations ofolfactory response were 0.8 and 54.1 s, respectively.The mean duration of the olfactory response of malesand females was

8.6

±

0.4

s (from 0.8 to 51.8;

n

= 222)and

8.9

±

0.7 s (from 1.4 to 54.1;

n

= 110), respectively.The duration of the olfactory response to various

excretions differed in males and females (Fig. 3). The

longest response was evoked by the perineal glandsecretion. The responses to other excretions, urine andfeces, were much shorter. Males sniffed urine longerthan feces, while females, conversely, sniffed urineshorter than feces.

Differences in the responses of the palm civet to theperineal gland secretion, urine, and feces were con-firmed in special experiments (Table 2) with pairs ofdifferent secretions of the same animal simultaneouslypresented to the recipient: the perineal gland secretionand urine, perineal gland secretion and feces, or urineand feces. No significant difference in the olfactoryresponse to the perineal gland secretion and urine of thesame animal was observed in both males and females.In contrast, significant differences were observed in theanimals of both sexes in their response to the perinealgland secretion and feces of the same individual. Theolfactory response to the simultaneously presentedurine and feces differed only in male palm civets, whichusually sniffed feces longer than urine; no significantdifference in the response of females was observed. Thepresented experimental data confirm the revealed dif-ferences in the duration of the palm civet olfactoryresponse to various types of their excretions.

Table 1.

Numbers of different forms of scent-marking behavior in male and female palm civets calculated per 1 h observation

Form of behaviorMales Females

pn M

±

m

δ

n M

±

m

δ

Dragging the perineal gland 6 5.6

±

1.5 3.69 2 5.3

±

0.1 0.07 >0.05

Rubbing the ear–neck area 6 2.3

±

0.7 1.74 2 1.9

±

0.2 0.21 >0.05

Heel rubbing 6 1.5

±

0.5 1.24 2 0.2

±

0.1 0.14 >0.05

Anal drag 6 0.1

±

0.1 0.13 2 0.2

±

0.2 0.21 >0.05

Urination 6 0.7

±

0.1 0.29 2 0.3

±

0.2 0.21 >0.05

Defecation 6 0.3

±

0.1 0.19 2 0.2

±

0.1 0.14 >0.05

Total 6 10.4

±

2.3 5.62 2 7.9

±

0.5 0.64 >0.05

1 2 3 4 5 6

0

20

40

60

80

100

51.448.6

54.845.2

88.2

11.8

33.3

66.7 70

30

60

40

MalesFemales

Fig. 2.

Proportions (%) of the forms of scent-marking be-havior displayed by male and female palm civets, relative tothe total number of displays by the animals of both sexes(figures in parentheses): (

1

) dragging the perineal gland(678), (

2

) rubbing the ear-neck area (260), (

3

) heel rubbing(144), (

4

) anal drag (12), (

5

) urination (66), and (

6

) defeca-tion (28).

5

10

15

20

25

0

Secretion Urine Feces

19.1

12.8

5.7 6.44.9

6.9

MalesFemales

Fig. 3.

Duration (

c

) of olfactory responses to perineal glandsecretion, urine, and feces in male and female palm civets.

c

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Information Content of Scent Marks of the Palm Civet

The data on the ability of the palm civet to obtaininformation about the species, sex, and familiarity ofanimals from their various scent marks after presenta-tion of two odor sources are given in Table 3. This abil-ity was estimated from the asymmetry of olfactoryresponse, i.e., its direction to one of the two simulta-neously presented odors and different durations of theirsniffing. The direction of the response was expressed by

the proportion of tests in which one of the two odorswas examined longer.

Perineal gland secretion.

The asymmetry in thepalm civet olfactory response to the perineal glandsecretion after presentation of this excretion of theirown and different species, male and female, as well asfamiliar and unfamiliar individuals was very pro-nounced. The palm civets reliably distinguished theperineal gland secretion of their own and different spe-

Table 2.

Direction of the palm civet olfactory response to various excretions and the mean difference (

M

±

m

) of sniffingduration for each pair of odors, s

Pair of odors

n

Directionof the response

M

±

m p

Males

Perineal gland secretion–urine 27 15 : 12 1.0

±

0.8 >0.05

Perineal gland secretion–feces 12 12 : 0 5.4

±

1.1 <0.01

Urine–feces 28 5 : 21 –1.2

±

0.4 <0.01

Females

Perineal gland secretion–urine 21 13 : 7 2.1

±

1.0 >0.05

Perineal gland secretion–feces 10 10 : 0 3.9

±

0.7 <0.01

Urine–feces 15 6 : 8 –0.4

±

1.5 >0.05

Table 3.

Direction of the palm civet olfactory response to various scent marks and the mean difference (M ± m) of sniffingduration for each pair of odors, s

Pair of odors n Directionof the response M ± m p

MalesPerineal gland secretion

Own species–different species 17 1 : 16 –2.1 ± 0.4 <0.01Male–female 25 4 : 21 –8.6 ± 2.6 <0.01Familiar male–unfamiliar male 13 0 : 13 –3.3 ± 1.0 <0.01

UrineOwn species–different species 12 0 : 12 –3.0 ± 0.6 <0.01Male–female 27 4 : 20 –3.3 ± 0.9 <0.01Familiar male–unfamiliar male 16 1 : 14 –0.8 ± 0.2 <0.01

FecesOwn species–different species 11 0 : 11 –4.1 ± 1.2 <0.01Male–female 21 8 : 13 –0.6 ± 0.5 >0.05Familiar male–unfamiliar male 13 4 : 6 –0.1 ± 0.3 >0.05

FemalesPerineal gland secretion

Own species–different species 10 0 : 10 –2.6 ± 0.6 <0.01Male–female 15 1 : 14 –6.3 ± 2.2 <0.01

UrineOwn species–different species 6 0 : 6 –2.8 ± 1.4 <0.01Male–female 14 8 : 6 –0.7 ± 0.7 >0.05

FecesOwn species–different species 6 0 : 6 –2.5 ± 0.6 <0.01Male–female 13 8 : 5 –0.8 ± 0.9 >0.05

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cies (in virtually all cases, both males and femalessniffed the secretion of other species longer), of differ-ent sexes (in most cases males and females sniffed thefemale secretion longer), as well as of familiar andunfamiliar individuals (the experiment was carried outon males only, and they sniffed the secretion of an unfa-miliar individual longer).

Urine. The asymmetry in the palm civet olfactoryresponse to the presentation of urine of their own anddifferent species, male and female, as well as familiarand unfamiliar individuals was also very pronounced.Both males and females reliably distinguished urine oftheir own and different species and sniffed urine of adifferent species longer in all cases. The response ofmale palm civets to urine of the animals of differentsexes significantly differed, and they sniffed the femaleurine longer; while no significant differences in thefemale response to urine of both sexes were observed.The palm civet response to urine of familiar and unfa-miliar individuals reliably differed (the experiment wascarried out on males only, and they sniffed the secretionof an unfamiliar individual longer).

Feces. Simultaneous presentation of two feces odorsrevealed the asymmetry in the palm civet olfactoryresponse in both males and females only in the case offeces of their own and different species. The animalsalways sniffed feces of a different species longer. Noreliable differences in the response of both sexes tofeces of males and females as well as familiar and unfa-miliar individuals (this experiment was carried out onmales only) have been revealed.

DISCUSSION

Our observations and experiments with palm civetsdemonstrate that the frequency of marking, olfactoryresponse, and information content differ for variousexcretion types. These tropical carnivorous mammalsleave the perineal gland secretion most frequently, theirolfactory response to it is the longest, and the informa-tion content of these gland secretion entirely comple-ments that of urine scent marks.

Well-developed specific skin glands can be found inmany representatives of viverrids, which demonstratevariable ways of leaving their secretion in the environ-ment, including those mediated by the perineal gland.We have also revealed different forms of scent-markingbehavior in the palm civet: dragging the perineal gland,rubbing the ear–neck area, heel rubbing, anal drag, uri-nation (including specific form of behavior resulting ina urine path on a substrate), and defecation. The data onthe palm civet behavior allow us to propose differentsignificance of the above-mentioned forms of behaviorfor mediated communication by scent marks of this ani-mal. Dragging the perineal gland has the leading role inmarking behavior of males and females; this behavioris most commonly observed in the palm civet. Leavingthe perineal gland secretion is the most important mark-

ing behavior in other viverrid species too (e.g., in gen-ets, small Indian civet, and binturong) (Kleiman, 1974;Roeder, 1978, 1980, 1983; Rozhnov, 1994; Xu andSheng, 1994). These data suggest a common pattern ofbehavior with respect to the role of the perineal glandsecretion in mediated communication by scent marks inviverrids.

Rubbing the ear–neck area has a considerable role inscent-marking behavior of both males and females.This area apparently contains hypertrophied skinglands, and such a behavior is aimed at leaving theirsecretion on the substrate. However, no morphologicaldata confirming this assumption are available.

In addition to these two main forms of scent-mark-ing behavior of the palm civet, heel rubbing and analdrag seem significant in males and females, respec-tively. Note that the soles of hind feet and the analregion of the palm civet have well-developed specificskin glands with a complex structure. They releasesecretions that apparently play a role in mediated com-munication by scent marks (Sokolov and Neklyudova,1988; Rozhnov et al., 1992); however this role stillremains unclear.

We also demonstrated that the scent-marking activ-ities of different types differ in male and female palmcivets. This difference is particularly pronounced forheel rubbing, which is much more common in males, aswell as for anal drag and urination, which are morecommon in females. The total scent-marking activityalso differed between males and females; males lefttheir scent marks more often. Similar data are knownfor other viverrid species, in particular, for genets dem-onstrating sex- as well as reproductive and social sta-tus-dependent scent-marking activity (Roeder, 1978).

Thus, leaving the perineal gland secretion is mostfrequently observed and, apparently, most importantform of scent marking behavior in mediated communi-cation by scent marks of the palm civet.

This is also confirmed by the duration analysis ofolfactory response of these animals to various excre-tions. The duration of the palm civet olfactory responseis quite variable; it differs in the animals of different sexand depends on the type and combination of the pre-sented excretions. Generally, the olfactory response ofthe palm civet to the perineal gland secretion waslonger. The duration of the olfactory response to urinewas longer in males than females, while the response tofeces was conversely longer in females than males.However, males in most cases sniffed feces longer thanurine in the experiments with simultaneous presenta-tion of these excretions. We believe that this can alsoconfirm different significance of urine and feces inmediated communication by scent marks of the palmcivet.

Korytin and Azbukina (1975, 1986) studied theolfactory response of mammals to various odor sourcesand demonstrated that its duration depends on manyfactors. In particular, they include animal species, sex,

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ROLES OF DIFFERENT TYPES OF EXCRETIONS IN MEDIATED COMMUNICATION 589

and age, as well as the nature of odor sources and phys-ical conditions (climatic and meteorological) of odorperception by the animal. The data obtained in ourstudy demonstrate the dependence of olfactoryresponse duration on sex of the recipient animal as wellas the type of sniffed excretions. It was shorter in malesand longer in females, although no significant differ-ences have been revealed. As concerns the dependenceon the type of excretion, we have observed the longestolfactory response in the experiments with the perinealgland secretion, while it was the shortest in the case offeces and urine for males and females, respectively.

Thus, the data on the duration of olfactory responseto various excretions in the palm civet confirm the pro-posed leading role of the perineal gland secretion inchemical communication between the animals of thisspecies.

Great significance of the perineal gland secretion aswell as urine in mediated communication by scentmarks of these tropical carnivorous mammals is alsoconfirmed by specific experiments when excretions ofdifferent types but from the same donor were presentedto the recipients animals in pairs: the perineal glandsecretion and urine, the perineal gland secretion andfeces, as well as urine and feces. Significant differencesin the olfactory response to the perineal gland secretionplus other secretion type have been revealed for ani-mals of both sexes; both males and females sniffed theperineal gland secretion longer in all cases.

The experiments on the information content of var-ious scent marks confirmed once again that the perinealgland secretion, urine, and feces have different signifi-cance in mediated communication by scent marks ofthe palm civet, with the perineal gland secretion play-ing the most significant role.

The palm civet receives all types of informationstudied in this work (species specificity, sex, and famil-iarity) from the odor of the perineal gland secretion.This also applies to urine marks. According to our data,the palm civet receives considerably less informationfrom feces odor—only species specificity of the scentmarks.

The observed likeness of information transmitted bythe perineal gland secretion and urine marks is quiteinteresting. Apparently, the scent marks of the perinealgland secretion are preserved for a longer period undervarious environmental influences and function as along-term source of information for the animals. As aresult, frequent renewal of the scent marks is notrequired, on the one hand, and reliability of the medi-ated communication by scent marks increases, on theother.

The data on different roles of various excretions areavailable for other carnivorous mammals. Genetsreceive considerably less information from feces odoras compared to other excretions, particularly, urine(Roeder, 1980). The European polecat and American

mink also receive more diverse information from urinethan feces odor (Sokolov and Rozhnov, 1983, 1989).

Thus, mediated communication by scent marks ofthe palm civet, a representative of tropical carnivorousmammals of the family Viverridae, has a number of fea-tures related mainly to the presence of the specific skingland. They include dragging the perineal gland, themost commonly observed scent marking behavior ofthe palm civet; the longest olfactory response of thepalm civet to this gland secretion, which reflects greatattention of the animals to this excretion; and the abilityof the palm civet to receive complete information aboutthe other animal (species, sex, and personality) fromthe odor of this secretion. These data suggest that theperineal gland was formed during the evolution to pro-vide for a more reliable mediated communication byscent marks.

ACKNOWLEDGMENTS

We are grateful to I.V. Korneev, director of the Len-ingrad Zoo (St. Petersburg), for providing an adultfemale palm civet for the experiments and to A.A. Pet-rin, T.N. Petrina, Sv.V. Naidenko, and O.O. Nikitina(Tchernogolovka Biological Station) for their help inthe experiments. This work was partially supported bythe Russian Foundation for Basic Research, project no.00-04-55001, and the Integratsiya Foundation for Spe-cial Research Programs, project no. A0084.

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