Comparative Ethology of the

Large-spotted Genet (Genetta tigrind)

and Some Related Viverrids

CHRISTEN M. WEMMER

m

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 239
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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 239

Comparative Ethology of the

Large-spotted Genet Genetta tigrina

and Some Related Viverrids

Christen M. Wemmer

SMITHSONIAN INSTITUTION PRESS

City of Washington

1977
ABSTRAC T

Wemmer, Christen M. Comparative Ethology of the Large-spotted Genet

(Genetta tigrina) and Some Related Viverrids. Smithsonian Contributions to

Zoology, number 239, 93 pages, 37 figures, 18 tables, 1977.—The study describes

and analyzes the behavior of the large-spotted genet (Genetta tigrina) in captivity, and treats selected aspects of behavior in eight viverrid genera occupying

five different subfamilies. The natural history of the genus Genetta is reviewed,

and the relationships of the living members of various subfamilies are discussed.

The behavior of solitary animals is treated by considering locomotion, stances,

and object- and self-oriented contact patterns. Scent-marking with glandular

products, urine, and feces is considered with reference to the communicative

properties of the marking behavior and the substance. Sonographic and contextual analyses are used to classify vocalizations and serve to establish homologies between the calls of several species. Further analyses of social behavior

in Genetta and Civettictis are based on encounters within all possible pairs of

animals. A small number of behavior patterns predominates during most interactions between animals of both species. The primary differences in the communication systems of these two species are probably related to semi-arboreal as o p

posed to terrestrial habits.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea

cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Wemmer, Christen M.

Comparative ethology of the large-spotted genet (Genetta tigrina) and some related viverrids.

(Smithsonian contributions to zoology ; no. 239)

Bibliography: p.

1. Genetta tigrina—Behavior. 2. Genetta—Behavior. 3. Viverridae—Behavior. I. Title. II. Series: Smithsonian Institution. Smithsonian contributions to zoology ; no. 239.

QL1.S54 no. 239 [QL737.C28] 591'.08s [599'.74422] 76-15195
Contents

Page

Introduction 1

Background 1

Intra-viverrid Affinities 2

Behavioral Classification and Description 4

Natural History of the Genus Genetta 5

Materials and Methods 6

Acknowledgments 9

Behavior of Solitary Animals 9

Primary Senses 9

Locomotion and Its Ontogeny 10

Shaking and Twitching Movements 16

Stationary Body Positions 17

Stretching Movements 18

Object-oriented Contact Patterns 20

Object-oriented Contact Patterns in Other Genera 23

Communicative Function of Object-oriented Contact Patterns in

Genetta 26

Self-oriented Contact Patterns 29

Self-oriented Contact Patterns in Other Genera 32

Ontogeny of Shaking, Stretching, and Self-directed Behaviors in

Genetta 32

Elimination 36

Communicative Functions of Elimination in Genetta 36

Some Intergeneric Comparisons and Derivational Schemes 37

Behavior of Associated Animals 42

Companion-oriented Locomotion 42

Companion-oriented Stances 43

Noncontact Body Movements 44

Companion-oriented Contact Patterns 47

Head 47

Limbs 49

Trunk 50

Compound Contact Patterns 50

Companion-oriented Contact Patterns in Other Genera 50

Vocalization 51

Vocalization in Other Genera 54

Analyses of Interaction 59

Materials and Methods 59

Results 60

Effects of the Encounter Introduction Schedule on Behavior in

Genetta 60

General Features of Encounters 61

Interaction through Sniffing and Head-darting: the Sniff-Dart Axis . . 74

Discussion 76

Vocalizations among Viverrids 76

Nonvocal Interaction in Genetta and Civettictis 79

Addendum 80

Appendices: Encounter Schedules and Statistical Procedures 81

Literature Cited 90

iii

Comparative Ethology of the

Large-spotted Genet {Genetta tigrina)

and Some Related Viverrids

Christen M. Wemmer

Introduction

BACKGROUND

The genus Genetta occupies the niche of the

small, semiarboreal, solitary, nocturnal, insectivorecarnivore. The dozen or so species comprising the

genus are structurally coherent, exploit nearly all

natural and cultivated habitats over most of Africa

and southwest Europe, and in many areas are a

numerically dominant small carnivore. This success

may be considered exceptional, for Genetta, like

Didelphis and Tupaiia, is believed to be a relatively

unmodified derivative of an ancestral form

(Gregory and Hellman, 1939; Colbert, 1961). The

view that the viverrines are the most structurally

conservative viverrids, however, rests on the premise

that tribosphenic molars, attenuated body form,

and relatively unrestricted rotatory ability of

manus and pes constitute features of the ancestral

morphotype (Gregory and Hellman, 1939; Colbert,

1961; and Romer, 1945). Since both fossil viverrines,

such as the Oligocene and Miocene genera Paleoprionodon and Stenoplesictis, and extant civets such

as Prionodon, Viverricula, Poiana, and Genetta

more or less share these features, the subfamily

Viverrinae is considered to be the most central and

basal viverrid group (Winge, 1941; Thenius and

Hofer, 1960; Gregory and Hellman, 1939; Colbert,

Christen M. Wemmer, Conservation and Research Center, National Zoological Park, Smithsonian Institution, Front Royal,

Virginia 22630.

1961; and Romer, 1945). Whether the lineage of

the more terrestrial civets {Viverricula, Viverra,

Civettictis) or the semiarboreal civets (Poiana,

Genetta, Prionodon) represents the least altered

derivative of the common ancestor is debatable, for

the fossil record is inadequate. What little is known

about Poiana richardsoni, though, suggests that a

semiarboreal habitus may more closely approximate

the ancestral viverrine condition.

Besides displaying more cranial resemblance to

Paleoprionodon than other extant genera (R. H.

Tedford, pers. comm.), Poiana is purported to

build leafy arboreal nests (Walker, 1968), a trait

interpretable as primitive by comparative carnivore

standards. Furthermore, Booth (1954) has interpreted Poiana and other West African species

having disparate distributions to be stabilized forest

forms that reached an evolutionary climax before

the appearance of the Dahomey Gap. Genetta's

capacity for radiation and dispersal into woodland

and savanna is a progressive feature that presumably emerged early in its lineage.

Contrary to the prevailing view of the genet, it is

herein acknowledged as a conservative and generalized solitary predator, but not as a model

antecedent of more specialized contemporary viverrids; rather, the genus displays a complex of morphological and behavioral characteristics some of

which are considered to be fundamental to the

evolution of more advanced or specialized forms.

With this perspective in mind, the present study

attempts (1) to establish qualitative and quantita1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tive behavioral characteristics, and to outline the

principle features of the communication system in

captive genets, and (2) to compare certain behaviors

of selected viverrids and interpret the results in

light of known taxonomic affinities.

INTRA-VIVERRID AFFINITIES

Because modern taxonomy strives to arrive at

evolutionary classification by delimiting taxa on

the basis of common ancestry and subsequent divergence (Mayr, 1968), behavioral evolution must ultimately derive from morphologically based appraisals

of systematic position. Though behavioral criteria

are often useful in elucidating relationship (Lorenz,

1953; Lawrence and Novick, 1963; and McKinney,

1965, 1966), their role in clarifying the affinities of

higher taxa (families, orders) is minor. Unfortunately, the Viverridae have received so little attention that it is questionable whether the existing systematic schemes serve the practical needs of a

sound classification. Many genera are rare or confined to remote tropical regions, resulting in a

scarcity of study animals as well as few preserved

specimens. In addition, a number of genera and

perhaps one subfamily exhibit character traits that

defy a simple scheme of filial relationship. While

the pretense of a solution might be made by referring only to the most recent synopses of taxa

(Anderson and Jones, 1967; Wenzel, 1972), it would

ignore the problematic status of several genera

dealt with herein.

Though a large number of viverrid species was

available to me for comparative study, it became

increasingly apparent, as it had to others (Diicker,

1971; Ewer, pers. comm.), that the mongooses (Herpestinae) exhibited a more uniform and distinctive

level of behavioral organization than other viverrids. Similar observations were made by Pocock

(1916b) and Gregory and Hellman (1939) who had

advised elevation of the mongooses to familial

status on the basis of their uniformity in foot,

dental, and glandular characteristics. This suggestion was subsequently endorsed by Wurster and

Benirschke (1968:374), who found that eight species

and seven genera of mongooses examined karyotvpicallv were "all nearly identical to one another

and without exception, they bear no satellited

marker chromosome." In addition, Ewer (1973),

noting that all four known herpestine genera had

horizontal oval pupils, stated that this structure

may be characteristic of the Herpestinae as a whole.

A vertical slit pupil is found in all the other known

viverrids with the exception of Paradoxurus whose

pupil is a horizontal slit (Table 1; cf. Walls in

Ewer, 1973). Lastly Radinsky (1971) has shown that

in 25 of the 36 living genera representing all subfamilies, Crytoprocta, the herpestines, and the galidiines all possess a well-developed cruciate sulcus,

while the structure is small or absent in viverrines,

paradoxurines, and hemigalines.

The existence of the Madagascan Galidiinae

accounts for some of the difficulty in distinguishing

the mongooses (Herpestinae) from the bulk of the

viverrids. Both cranially and in gross body form the

subfamily shows a strong affinity with the herpestines, but unlike the latter they possess the carnivore

satellite chromosome (Table 1). Carlsson (1910)

showed that Galidia elegans shared 14 anatomical features with the other Malagasy civets (including Ciyptoprocta and Eupleres), 10 with the

mongooses, and 7 with the viverrines, but he concluded that relegation of the Galidiinae to either

the viverrids or herpestids would obscure its position as a stem form that differentiated prior to the

evolution of either the herpestine or viverrine grade

of organization. Gregory and Hellman's decision

to place the galidiines within the Viverridae was

consistent with their implicit belief that the rate

of evolution in viverrids (particularly viverrines)

has been much slower than in herpestids.

Though the fossa (Cryptoprocta ferox) of Madagascar is distinctive at the subfamilial level, it too

displays features of conflicting affinity. Although

Pocock (1916a) gave the fossa familial status,

Gregory and Hellman placed it as a subfamily

within the Felidae, believing it to be an excellent

structural link between the primitive Viverrinae

and the Felidae. This opinion was predicated on

skeletal similarities. While its auditory region and

dentition resemble the viverrines and felids respectively, the presence of an anal pouch (and the

absence of the perineal gland) prompted Mivart to

ally this species with the mongooses (in Gregory

and Hellman, 1939). Karyotypically, however, the

fossa has much in common with the paradoxurines, hemigalines, and galidiines (Wurster and

Benirschke, 1968). Thus, without interpreting the

fossa's affinity, it appears to be an early viverrid
NUMBER 239

TABLE 1.—Comparisons of major characteristics among viverrid subfamilies and

genera (- = absence within taxon, + = presence within taxon but not necessarily universal, ± = present in some members of taxon, when no character

alternatives listed)

Characteristics t - T3

Skin Glands

Perineal pouch —

Preputial pcoich...

Anal gland ,

Anal pouch

Feet

Plantigrade

Sub-digitigrade...

Digitigrade

Digit reduction...

Retractile claws..

Teeth

Sectorial

Tuberculate

Molar reduction...

Food Habits

Carnivorous

Omnivorous ,

Piscivorous

Insectivorous

Myrmecophagic

Coloration

Spotted

Striped

Banded

Uniform

Misc. Features

Marker chromosome

Vertical pupil

Distribution

Africa

Madagascar

Asia

Europe

survivor displaying convergent or possibly archaic

incipient felid features.

The African palm civet, Nandinia binotata, resembles the paradoxurines in both its gross body

form and its skull. On the basis of its more

trenchant carnassials and cartilaginous auditory

bulla Gregory and Hellman surmised, however,

that this civet branched off from the paradoxurine
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

stem prior to frugivorous specialization. The existence of the subossified or cartilaginous entotympanic bone of the auditory bulla is generally

regarded as a secondarily derived feature rather

than a vestige of a miacid ancestry. The extraordinary nature of this structure and the presence

of a preputial scent gland prompted Pocock (1915b)

to separate Nandinia as a family. Ironically, the

species strongly resembles the viverrines in its

possession of a subtelocentric marker and a small

acrocentric Y-chromosome (Wurster and Benirschke,

1968).

The affinity of Asian linsangs (Prionodon) to the

viverrines resembles that of Nandinia to the paradoxurines. While sharing features of body form

with Genetta and karyotype with the the viverrines

in general, Prionodon lacks the metatarsal pads, the

perineal gland, and the second upper molar. These

differences were interpreted by Pocock (1915a) as

specializations justifying a distinct subfamilial status. Similarly, Gregory and Hellman relegated the

linsang to a viverrine tribe (Prionodontini).

The fanaloka (Fossa fossa) in its possession of

vestigial metatarsal pads and a genetlike hair pattern is superficially somewhat viverrine; karyologically and cranially it is distinctly hemigaline, while

in gross body form it is remarkably foxlike. It is

probably an early Asian hemigaline waif to Madagascar that has convergently acquired a foxlike

habitus.

BEHAVIORAL CLASSIFICATION AND DESCRIPTION

Ethological studies have traditionally relied upon

both functional and motivational criteria in categorizing a species' behavioral repertoire. Many

behavior patterns, however, can be assigned to

several categories, others can be accommodated by

none, and new criteria are invoked in order to

categorize special cases (e.g., displacement activities).

To design an objective, systematic scheme with

mutually exclusive categories is both difficult and

overreductionistic, because in most animals a large

number of biological roles must be served by a

number of movement patterns restricted by the

mechanical and physiological properties of the species' body structure. As much as functional and

motivational considerations may enhance a descriptive work, an accurate understanding of these

factors requires both experimental validation and

a knowledge of the species in its natural realm.

Because these labors exceed the requirements of

faithful description, it seems more objective to

classify behavior on observable and measurable

characteristics of form, rather than on deduction

about motivational state, for instance. It will be

noticed that though some new terms are used for

descriptive categories, implicitly and explicitly functional terms, such as grooming, scent-marking, and

play, are retained within the descriptions.

Behavioral description has for the most part been

an empirical process; the events ethologists recognize and interpret as true behavioral units (action

patterns) are assumed to represent fuctionally meaningful acts unabstracted by the observer's conceptual

processes. But in mammals, at least, it is questionable whether the discrete phasic properties of action

continua accurately represent all the meaningful

dimensions of animal conduct. Symposia titles such

as "The Search for Natural Units of Behavior"

 x

convey the distinct impression that (1) recognized

behavioral units may be abstractions rather than

true or complete representations of natural phenomena, (2) the types of units currently used have

limited analytical utility, and (3) the elucidation of

"natural units" will hopefully permit more insight

into the meaning of animal behavior. With the

exception of locomotion, which received early and

exacting description by virtue of historical and

esthetic interest (Muybridge, 1957), most forms of

mammalian behavior have not received the analytical descriptions exemplified by the Baerends' (1950)

cichlid work or Dane, et al. (1959) and McKinney's

(1965, 1966) studies on duck behavior. In particular,

there have been few critical attempts to establish

either indices of variability or typical intensity

(Morris, 1957) of mammalian displays (Le Boeuf

and Peterson, 1969, is one exception).

The Eshkol-Wachmann Movement Notation

Method (Golani, 1969), which permits fine-grain

temporal and spatial resolution of behavior, has

remained an unexploited descriptive tool. With

rigorously objective description, the method provides a valuable alternative to the interpretive

methods that have prevailed up to now, though

from the standpoint of comparative ethology it

'A lecture presented by S. A. Altmann at a symposium

titled "Analytic Problems in the Study of Animal Behavior,"

67th Annual Meeting of the American Association for the

Advancement of Science (1970).
NUMBER 239

requires considerable time. Useful descriptions can

be achieved more expediently. Not infrequently,

however, descriptions are made in such general

terms that they have limited utility: it is often

necessary to know the postural and locomotory

details as well as the basic mechanics of a motor

pattern in order to appraise it comparatively. For

this reason, photography is often as integral to the

analysis and description of body motion as sonograms are to the description of sound.

The classification used in this study is a phenetic

one; patterns are lumped together on the basis of

overall similarity in form. The system replaces

functional and motivational pigeonholing. For

example, the patterns usually ascribed to the maintenance (or grooming) behavior category are herein

subdivided into classes of self- and object-oriented

contact patterns. The latter category also encompasses behaviors that are otherwise classed as comfort movements (e.g., body rubbing) and scentmarking. As Mayr (1968) has remarked, no

classification is absolute because the structuring of

character sets always involves a subjective element.

The present system does not claim to be consistent

in its criteria. Although the orientation of the

animal's body parts to one another and to the

environment is a primary consideration, certain

behaviors (e.g., vocalization) do not fit the scheme.

The utility of the system, however, lies in bringing

to attention certain elemental similarities in behavioral form.

NATURAL HISTORY OF THE GENUS Genetta

The separation of the genus Genetta into three

subgenera is based upon characters of the skull,

teeth, and foot pads. The majority of species are

placed within the subgenus Genetta and these are

distinguished by the patterning of spots, spinal

stripes, and tail rings (in addition to cranial and

dental criteria). Forest species are on the whole

darker in color and have more spots than savanna

forms (Matschie, 1902). Genetta victoriae appears

to be a somewhat aberrant species tending towards

large size and terrestriality. Its coloration shows

considerable resemblance to that of the African

civet (Civettictis civetta) with which it is sympatric

(Thomas, 1901).

Members of the subgenus Pseudogenetta (villiersi

and abyssinica) are characterized by relatively small

size and the lack of hair in the depression between

the interdigital and metacarpal pads. Paragenetta,

represented by one species (johnstoni), is known

only from skulls. The delicate dental structure of

this species suggests its diet may consist largely of

arthropods.

Genets occupy nearly all African habitats, but the

range of habitat tolerance varies between species.

Genetta victoriae, servalina, angolensis, johnstoni,

and mossambica are primarily forest dwellers,

while G. pardina occurs in forest-savanna ecotone,

woodland, and Guinea savanna. Drier nonforest

areas are occupied by G. genetta, tigrina (sensu

latu), villiersi, and abyssinica (Bigalke, 1968;

Coetzee, in press). Although Dorst and Dandelot

(1970) state that G. villiersi inhabits forests, a large

number of this species in the National Museum of

Natural History was collected from Sudan and

Guinea savannas in Ivory Coast and Upper Volta.

At least in Botswana, G. genetta penetrates considerably drier areas than tigrina, though both

species are often found together (R. H. N. Smithers,

pers. comm.). Displaying a tolerance for habitats

ranging from low forest to woodland and savanna,

G. tigrina has been collected from elevations as

high as 2560 m (8500 ft) on the slopes of Mt. Kenya

(Hollister, 1918). It often appears to be a "culturefollower," inhabiting the vicinity of villages and

cultivated land (Rahm, 1966).

There appears to be a distinct sexual dimorphism

as to size in some populations of G. tigrina (i.e.,

stuhlmanni, pers. observ.), but data are inadequate

to make conclusive statements for other species.

J. C. De Meneses Cabral (in Coetzee, in press) has

found that, in areas of low rainfall, skull length in

G. rubiginosa (= G. t. rubiginosa, sensu lato) varies

from an average of 87.5 mm in females to 90 mm in

males, while in high rainfall areas skulls of both

sexes average 90 mm. Tables of the genet cranial

measurements in Allen (1924) suggest sexual differences in skull dimensions may exist in G. victoriae

and servalina as well.

Melanistic genets are sometimes seen in museum

collections and zoos (Sanderson, 1940; Maliniak,

pers. comm.). The natural frequency of the melanistic morph in the wild is unknown. At the National Zoo a wild-type female mated by a melano

produced two wild-type offspring (G. Maliniak, pers.

comm.).

All known species appear to be nocturnal and
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

solitary, though pairs and family groups are occasionally seen. Evidence at present indicates that

the most complex social unit is the mother family;

its integrity is probably maintained only until the

young are weaned. Ducker's (1957) observation of

a parturient female's antagonism towards its male

cage mate suggests that the sexes remain separate

during the rearing of young. Verheyen's (1951)

statement that the male participates in providing

food for the young is doubtful.

The exact nature of the social system, however,

is unknown. Presumably a land tenure system exists,

but information is sparse regarding activity in and

defense of the home area. In the course of capture

and release work with genets inclined toward

poultry thievery, Carpenter (1970) recaptured four

of six female G. rubiginosa ( = G. t. rubiginosa sensu lato) at their original capture points within days

after release from as far away as 20 miles. Six of 11

males were taken again at poultry runs, but none

was recaptured at the original trapping site. He

attributes the fact that most poultry thieves are

males to their greater mobility, and suggests that

females, unlike males, may be territorial. Cobb

(1970), on the other hand, found that in Turkana,

Kenya genets frequently crossed paths; he is of the

opinion that in this area, at least, genets are not

strictly territorial. Locus-specific defecation, a trait

often correlated with the territorial syndrome, also

characterizes G. tigrina (Verheyen, 1951; Rahm,

1966; and Carpenter, 1970). The tentative and

somewhat contradictory nature of all these findings

clearly indicates that more work must be done to

elucidate this aspect of the genet's natural history.

Breeding in G. genetta and G. tigrina (sensu

lato) occurs during the wet seasons in both Kenya

(Taylor, 1969) and Botswana (Smithers, pers.

comm.). Taylor (1969) collected lactating and pregnant females in May and from September to December, and believes the timing of reproduction during

these rainy periods to be linked with an abundance

of insects. Pregnant females from Botswana were

also taken during the warmer wet period from

October to February (Smithers, pers. comm.).

Whether wild females breed during both wet seasons is uncertain; however, a breeding pair at the

National Zoological Park has regularly produced

a litter of two in April or May and again in July

or August, for the past six years. Females have two

pairs of inguinal mammae, and the number of

young born in the wild varies from two to four

(Hollister, 1918; Shortridge, 1934; Ansell, 1960;

Copley, 1950; Roberts, 1951; and Verheyen, 1951).

The types of resting, parturition, and rearing

sites are probably influenced by habitat. Holes in

trees, in fallen logs, and burrows in kopjes and

cliff areas seem to be usual locations (Shortridge,

1934). Rowe-Rowe (1970) located three young G.

genetta in a disused crow's or hawk's nest about 6

m above ground.

Feeding appears to be opportunistic with a strong

predisposition for animal matter. In a sample of

78 G. genetta from Botswana Smithers (pers. comm.)

found that the three animal groups represented

most frequently in stomachs were (in decreasing

order) rodents, Orthoptera, and scorpions; in a

sample of 30 G. tigrina from Botswana, rodents

ranked first, followed by Coleoptera and Orthoptera. Cobb (1970) reported that on the basis of

stomach contents, fecal samples, and observation,

the genets of Turkana, northern Kenya fed primarily on birds which roost in the dominant shrub,

Salvador a persica.

Nothing is known about the natural predators of

genets; they are presumably preyed upon by larger

carnivores and birds of prey, and are sought by

humans for food and fur. According to Dekeyser

(1956), G. villiersi is killed by the Bassari of Itiolo,

Senegal, for its tail, used in ceremonial rituals.

Genetta's climbing ability probably provides an

important niche parameter, not only in facilitating

escape into arboreal refuge, but also in making

accessible sources of food and cover that are denied

to strictly terrestrial animals.

MATERIALS AND METHODS

Table 2 presents personal history information

concerning the 11 genets on which the results of this

study are based. Eight of these animals were born

in the Small Mammal House at the National Zoological Park, Washington, D.C. For convenience

they are referred to in the text by the following

abbreviations: K (Kitzi), C (Clawdina), I (Ivy), Mo

(Monika), M (Marvin), O (Oscar), E (Elsie), Fr

(Froda), and F (Felix). The origin of the parental

pair is unknown, though comparison of skins and

skulls of the previous offspring of this pair with

specimens at the Smithsonian Institution reveals

similarity to Genetta tigrina stuhlmanni, a sub-
NUMBER 239

TABLE 2.—Caging history (cage locations, cagemate(s), periods of confinement,

cage dimensions in meters) of study animals (Genetta tigrina)

Subject

Parent (P)

Parent (P)

Kitzi (K)

Clawdina (C)

Ivy (I)

Monika (Mo)

Marvin (M)1

Oscar (0)

Elsie (E)'

Froda (Fr)

Felix (F)2

Sex

0*

?

o"

5

?

?

i

<S

?

?

0*

Date of

birth

Before 1966

Before 1966

22 Apr 67

22 Jul 67

22 Jul 67

01 Feb 68

01 May 68

24 Apr 69

14 May 70

05 Aug 70

? Jan 70

University of Maryland

C.I Nov 1967—Jun

1.37 x 2.43 x

K,I Nov 1967—Jun

1.37 x 2.43 x

C,K Nov 1967—Jun

1.37 x 2.43 x

M Aug 1968—Dec

1.37 x 2.43 x

Dec 1968—Jun

1.37 x 2.43 x

Mo Aug 1968—Dec

1.37 x 2.43 x

— Oct 1969—Jun

1.37 x 2.43 x

- Jul 1969—Jun

1.21 x 1.21 x

1970

2.43

1970

2.43

1970

2.43

1968

2.43

1970

2.43

1968

2.43

1970

2.43

1970

2.43

9 P

<f P

C

K

F

E

0

~

I

National Zoo

Duration of study

Duration of study

Sep 1970—Jun

1.90 x 1.90 x

Sep 1970—Jun

1.90 x 1.90 x

Sep 1970—Jun

1.90 x 1.90 x

Sep 1970—Jun

.91 x .91 x

Sep 1970--Jun

1.90 x 1.90 x

Sep 1970—Jun

1.90 x 1.90 x

Sep 1970—Jun

.71 x 1.21 x

Sep 1970—Jun

1.90 x 1.90 x

1971

2.03

1971

2.03

1971

2.03

1971

1.21

1971

2.03

1971

2.03

1971

1.21

1971

2.03

* Died during anesthesia in January, 1969.

2

 Wild-born near Bangui, Central African Republic.

species inhabiting northwestern Tanzania (Coetzee,

in press). In addition, one male Genetta rubiginosa

(— G. t. rubiginosa) from the vicinity of Bangui,

Central African Republic, was obtained as a hand

reared specimen from Mrs. C. Doscher.

Kittens were always taken from their mother on

the day of birth or the day after and were confined

to a box containing a heating pad and a blanket

until they were able to move about by themselves.

The first three hand reared genets had unrestricted

freedom in our house; subsequent animals were

confined to one room for most of the day. Their

diet consisted of warm Esbilac (Borden Company)

mixed with water at recommended proportions and

fed from a graduated (240 cc) nursing bottle designed for puppies (Evenflo Company). On two

occasions this diet produced repeated vomiting that

was remedied by substituting cow's milk mixed with

egg yolk. Weaning was initiated by feeding morsels

of raw meat from the nursing bottle nipple. Infants

were triggered to urinate and defecate by massaging

the perineum with a moist paper towel.

A couple of months after weaning, the hand

reared genets were transferred to a laboratory at

the University of Maryland. As the windows were

open most of the year, to some extent the animals

experienced local temperatures and humidity, except during thunder showers, cold spells, and winter
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE I.—Genet cages used at the University of Maryland:

A, a cage for single animals (0.71 X 1.21 X 1.21 m); B, a large

cage for two or three animals (1.37 X 2.43 X 2.43 m).

when the windows were closed and the room maintained at about 13°-19° C (55°-60° F). During the

summer when temperatures reached 27° C (80° F)

an electric fan circulated the air.

Cages of various sizes were used (Table 2; Figure

1A,B). All cages were provided with one or more

elevated nest boxes, vertical, horizontal, and diagonal branches of various diameters, and one or two

pans containing corncob litter (Sanicel) that served

as elimination sites for urination and defecation.

Diet consisted of chunk horsemeat with a vitamin

additive (Pervinal, Thayer Laboratories) obtained

from the National Zoological Park, supplemented

with rats and mice, and occasionally hens' eggs,

chicken embryos, chicks, and frogs. In the spring

of 1970, all the animals in my care were moved to

the Division of Scientific Research at the National

Zoological Park (Table 2) and were installed in

their permanent cages there by September of that

year. Maintenance was thereafter under the supervision of Mr. Gene Maliniak, head keeper for the

research collection.

Individuals of the following viverrid species were

also studied: a pair of Civettictis civetta (African

civet), two males and a female Nandinia binotata

(African palm civet), a pair of Paradoxurus hermaphroditus (Asian palm civet), a male Paguma larvata (Formosan masked civet), a pair of Arctictis

binturong (binturong), a male and two female

Hemigalus hartwicki (banded palm civet), two

pairs of Fossa fossa (fanaloka), three male Galidia

elegans (Malagasy ring-tailed mongoose), and a

male Cryptoprocta ferox (fossa). Animals were

observed in the National Zoo's Small Mammal

House sporadically in the evenings of 1967 and

1968, regularly during the early morning in June,

July, and August of 1968, and sporadically in the

afternoon (at feeding time) from 1968 to 1971. I

was also able to personally maintain several of these

animals for closer observation. A male Nandinia,

procured in 1967 from Kinshasa at about the age

of six weeks, was kept at home and later at the

University of Maryland until 1970. An adult female

Nandinia was also held at the University from 1968

to 1970, as was the male Paguma. In addition, a

male Fossa born in 1969 was hand reared at home

from the day of birth, and then held at the University until 1970. The male Civettictis, obtained as a

young adult in 1969, was also held for a year at

the University and then transferred to the National

Zoo when the young female was received. (These

animals were descendents of stock from Sierra Leone
NUMBER 239

(Mallinson, 1969).) In addition to the standard

meat diet, the Civettictis, Nandina, and Paguma in

my care were fed fruits, such as melon, strawberries,

apples, pears, oranges, grapes, and bananas.

Data for behavioral description were recorded in

writing, on tape recorders, on moving charts, and

on still and cine film. Equipment included a Uher

tape recorder (4000 Report-L) and microphone

(Model M-514), an Esterline Angus Operations Recorder, Praktica and Pentacon 35 mm SLR cameras,

a Pentacon 6 cm2 SLR camera, and Bolex P-4 8 mm

and H-16 Rex Reflex cine cameras. Drawings were

made from tracings of film projected on a Cabin

Hilldem 8 mm viewer and an LW photo-optical

Data Analyzer (Model 224-A). Figures and descriptions are based in part upon selections from a total

of 443.9 m (1400 ft) of 16 mm and 243.9 m (800 ft)

of 8 mm film. Sound spectrograms of tape recordings were made using a Kay Electronic Sonograph

(Model 6061 A) (85-8000 cps spectrum analyzer)

and a Kay Electronic Missile Data-Reduction

Spectrograph. Further details of method are described under specific chapter headings.

ACKNOWLEDGMENTS

Many people have contributed to different phases

of this study and to all of them I am grateful. I

especially thank James Murtaugh, Robert Cox,

Leon Davies, Robert Horwich, Murray Itzkowitz,

Richard Jachowski, George McKay, and Nicholas

Smythe for their contributions in materials, efforts,

thought, and companionship. I am grateful to

Larry Collins for his fellowship in work and for

his efforts in obtaining the facilities for genets at

the National Zoological Park; to Wyotta Holden

for her invaluable secretarial expertise and good

natured tolerance; to Mary McComas and Marleigh

Hartman for typing the final revision; to Gene

Maliniak for his care of the animals; and to

R. H. N. Smithers for the unpublished data he

kindly placed at my disposal. I thank Douglass

Morse and Wolfgang Schleidt for generously extending facilities, discussion, and enthusiasm during

the course of the study. My dissertation advisor,

John Eisenberg, must be credited with making the

study possible and for creating the stimulating and

unique research atmosphere at the National Zoo.

I extend my sincere appreciation to the University

of Maryland for graduate research financial assistance, without which the work could not have been

done; and to the Smithsonian Institution for

electing me a research associate while this report

was being prepared. Finally, I thank my wife for

her dedication and help in hand rearing eight

genets and for her goodwill to the animals that

lived with us.

Behavior of Solitary Animals

The following descriptions encompass behavior

patterns that appear in a variety of contexts, social

ones not excluded. The term "solitary," then, is in

a sense inadequate, though it indicates that these

behaviors do not require the presence of a conspecific as a source of stimulation or as a social

referent. Many of the patterns appear to be released

by such generalized stimulus properties that their

occurrence is not limited to a specific context,

though most of them appear predominantly in

certain circumstances. Distinctions in motivation

and function clearly exist, and many patterns have

a direct or indirect role in social communication.

The difference between these "solitary" behaviors

and those whose occurrence is more exclusively

linked to an association with other animals is one

of degree rather than kind.

PRIMARY SENSES

EYES.—Genetta tigrina has moderately large semifrontal eyes with vertically elliptical pupils. The

shape of the pupil varies from a narrow linelike

slit in bright light to a broad oval in dim light. The

color of the iris approximates that of the pelage;

at the time the eyes open (10-14 days) it is a dark

grayish brown that matches the pelage. During development the color of the eye fades to a medium

brown, but with a slightly more reddish hue than

the fur. Though the eyeballs can move within the

socket, their mobility is limited, and animals rely

heavily on head movement even to maintain visual

fixation of objects moving over small distances. Although the eyelids usually blink synchronously, it is

not uncommon to see a resting animal blink only

one. Diicker (1965) concluded that G. tigrina was

color blind when study animals were unable to distinguish between colored cards and gray ones

matched for brightness.

EARS.—The ear pinna of Genetta, like that of
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

other viverrids, is of the basic aeluroid type

described by Pocock (1916b) and Mohr (1952). But

it differs from other genera, with the exception of

Hemigalus, by having a finer layer of hair internally

and externally, and in being slightly larger though

lighter in build. The mobility of the pinna and the

relationship between ear motion and the reception

of different sounds merits special study. The orbits

can be moved through a horizontal arc of about

80° from a frontal to a lateral position; they can

also be deflected in a vertical plane to approximately the same extent. The orientation of the

pinna confers only slight changes in the size and

shape of the orbit when the ear is erect. The size

of the orbit, however, is greatly reduced when the

pinna is folded backwards, a position achieved

when the posterior edge is folded immediately

above the tragus and the anterior edge is bent at

the base. Two types of ear movement are easily

discerned. An ear may twitch once or several times

in rapid succession in response to loud sounds or

to tactile stimuli of the pinna or surrounding hair.

The second type of movement consists of alternating, asynchronous, horizontal deflections of the

pinnae accompanied by visual orientation toward

a sound and/or movement (Preyer's reflex: Nuboer,

1959).

In the absence of a visual target, the head often

scans from side to side in the direction of the

sound, gradually narrowing its arc. Alternating ear

movements of diminishing amplitude accompany

the head scanning, and when the head is motionless, the ears are often jerked alternately and sporadically, but with minimal amplitude. This movement appears to be an important means of

localizing unknown sources of sound, but the exact

mechanism remains to be elucidated. Certainly this

behavior is more pronounced in the genet than in

the other species.

NOSE.—The nose is important for both olfactory

and tactile perception. The thin mucous film of the

rhinarium (Figure 15F) probably facilitates olfaction by providing a continuous solvent interphase

from the external to internal nares. Also wind

direction may possibly be detected by differential

evaporation and/or cooling of the rhinarial surfaces.

Scent perception requires air passage across the

nasal epithelium, and apparently this is almost

always accomplished by accelerated, low amplitude

respiration. Though smelling (or sniffing) is often

a distinctive activity, it is sometimes difficult to

separate from respiration. In newborn genets, for

instance, the respiratory rhythm is often sporadic;

periods of rapid and shallow breathing alternate

with slow, deep breathing. In a six-day-old animal

(K), asleep after feeding, the number of respiratory

cycles varied from 27 to 41 per minute (x = 34.6,

N = 8).2 Audible sniffing was first noticed at the

age of two weeks (K, O, and E; N = 3). In its

mildest form it involved rapid but slight expansion

and contraction of the belly and rib cage and slight

changes in nostril shape. Intense sniffing is audible

and an extension and retraction of the mystacial

vibrissae in apparent synchrony with inspiration

and expiration can be seen. Sniffing is frequently

accompanied by a slight head-bobbing. While

smelling, the animal's body is stationary and the

head elevated but without an orientation to a

specific scent source (testing the air). On the other

hand, close range smelling of a specific object or

area often involves contact and usually occurs as

the animal walks.

LOCOMOTION AND ITS ONTOGENY

BODY FORM.—Genetta's lithe and versatile movement in trees and on the ground is a function of its

coordination and the mechanical properties of its

skeleton and muscles. Its attenuated axial skeleton

conforms to the usual carnivore pattern of 20 thoracolumbar vertebrae (Flower, 1966), but the 29

caudal vertebrae are excessive and by comparative

standards contribute to the tail's considerable

length (Table 3). Though the tail aids in adjusting

the body's center of gravity, its relative importance

to different species for arboreal locomotion is not

apparent from inferences of climbing proclivity

based on habitat selection. Table 3 shows, for

example, that the body proportions of the arboreal

Nandinia and Poiana are more closely approximated by those of G. genetta and G. villiersi, both

of which usually inhabit savanna. Assuming the

measurement to be reliable and the differences

between species adaptive, it appears that as a presumptive arboreal adaptation greater relative tail

length is not necessarily indicative of species inhabiting more woody habitat. Additional biogeographic

1

 Based on visual detection of rib cage movement; one to

three minutes interrupted each successive observation minute.
NUMBER 239 11

TABLE 3.—Some mean body measurements, proportions, sources and origins of several species of

Genetta and allied genera (top group of species from savannah, middle group from woodland

and forest, bottom group from forest; all measurements for specimens in National Museum of

Natural History, Smithsonian Institution, taken from specimen labels; measurements given in

millimeters)

Species

G. villiersi

(S. genetta

G^. tigrina

G_. pardina

G. servalina

G. victoriae

Nandinia

binotata

Poiana

richardsoni

9

9

49

9

<t

9

<f

9

<t

S

rf

9

<f

9

J9

<t

9

tf

Source

USNM

USNM

and

Hoi lister,

1918

USNM

and

Hoi lister,

1918

USNM

Allen,

1924

Allen,

1924

Allen,

1924

USNM

Allen,

1924

Allen,

1924

Origin

Senegal,

Upper Volta

Nigeria,

Kenya,

Senegal,

Upper Volta

Kenya,

Uganda

Ghana,

Ivory Coast

Congo

Congo

Congo

Ivory Coast,

Togo

Congo

Congo

Number

16

19

35

10

13

23

24

11

35

4

6

10

9

4

8

10

13

5

11

7

18

7

11

1

Head

and

body

427.6

409.2

417.6

474.7

474.1

474.3

463.5

446.2

458.3

510.0

507.3

508.7

455.0

457.0

489.0

485.0

573.0

573.0

467.3

464.5

466.2

510.0

480.0

332.0

Measurement

body parts

Tail

413.8

400.3

406.5

471.2

459.3

463.5

418.4

401.6

413.1

418.7

416.8

417.6

470.0*

406.0

475.0

446.0

444.0

448.0

498.9

529.2

510.7

570.0

544.0

380.0

of

Hind

foot

79.7

77.8

78.7

88.0

86.3

87.1

81.4

73.5

80.5

91.5

91.3

91.6

86.0

85.0

92.5

88.7

101.0

100.0

88.7

38.8

88.7

92.0

88.0

64.0

Percent

and body

Tail

96.1

97.8

97.3

99.2

96.2

97.7

90.2

90.0

90.1

32.9

82.1

82.1

103.5

88.9

97.1

91.9

77.4

78.1

106.7

113.9

109.5

111.7

113.3

114.4

of head

length

Hind

foot

18.6

19.0

18.8

13.5

18.3

18.3

17.5

17.5

17.5

17.8

18.9

13.0

18.9

13.6

18.9

18.3

17.6

17.4

18.9

19.1

19.2

18.3

18.3

19.2

* This figure appears to be an error: if the value was a misprint of 407.0, the corrected proportion of

tail to head and body, 89.4% , would be in accordance with the other values. Furthermore, the male

servalina specimen pictured in Allen (1924: Plate XIV, Figure 1) appears to have a tail shorter than

its head and body.

factors may be operative; indeed, longer tails may

be thermally adaptive in warmer, low rainfall areas

(Allen's rule: Mayr, 1963). Of the four woodland

and forest species of Genetta listed in Table 3, all but

servalina have relatively short tails and G. victoriae,

the largest genet, has the smallest feet.

As the legs are moderately short, the resulting

body carriage and center of gravity is low. An

ample degree of limb abduction is possible, particularly in the hindlegs. The manus and pes have

retractile claws and they are also capable of relatively unrestricted pronation and supination. As
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

noted by Taylor (1970), Genetta's manus is palmigrade and its pes semi-digitigrade. Foot pads are

smooth and somewhat edematous; in addition to

apical and interdigital pads, the metacarpus and

metatarsus are covered with medium-sized bipartite

pads. Genetta's foot pads have relatively less surface

area than Nandinia's, but more than Civettictis

(Figure 2). Climbing ability is proportional to the

extent of foot pad development. Figure 2 depicts a

series of feet with decreasing arboreal specialization.

With regard to the ontogeny of locomotion,

voluntary movement following birth is minimal,

though infants are usually capable of crawling by

the age of one or two days. The distinctive pattern

of movement at this time consists of a slow and

rhythmic, alternating lateral flexion of the neck

and anterior trunk. As the head sweeps from side

to side the philtrum and chin brush along the

substrate. The legs, unable to support the body,

are extended to the side with the soles of the feet

down; forward progress is achieved by pulling

movements of the forelimbs. Until four or five days

of age this locomotion produces a curving or circular path of movement. As animals begin to support their forebodies when walking, the path

straightens, and head-sweeping becomes both less

frequent and stereotyped. This movement of the

FIGURE 2.—Pad structure of the manus (upper row) and pes

in four viverrids: A, the African palm civet (Nandinia binotata); B, the large-spotted genet (Genetta tigrina); c, the giant

forest genet {Genetta victoriae); and D, the African civet

{Civettictis civetta). (Not to scale.)

head and forebody, appropriately termed "Kreisbogenschlagen" by Frank (in Ducker, 1965), apparently aids in tactile localization of the nipple, the

mother, and siblings.

At about the age of one week, young genets are

capable of supporting themselves and walking for

brief periods on the fore- and hindlegs, though the

latter tend to drag. On successive days, as the legs

assume a more adducted position under the body,

coordination improves, and the time required for

each stride decreases. The claws apparently cannot

be retracted for the first 8-14 days and easily snag

on irregular surfaces. Progression across a smooth

woolen blanket, for example, is greatly encumbered,

but on sloping and vertical surfaces the claws clearly

assist clinging (Figure 15A).

Even before infant genets are able to walk they

manifest a pronounced facility for maintaining

partially suspended and precarious positions. When

suspended by the head or neck, the tail and hindquarters execute a twirling motion common to

young of many species of mammals (Horwich, 1972).

If suspended by the tail, the forelimbs extend, but

the hindlimbs often flex with one clasping the other

against the belly. The latter behavior was not seen

on the week following birth, but was subsequently

performed regularly until the animals had developed the strength and coordination at about 3-4

months of age to right themselves by flexing their

torsos and climbing up their tails.

Once walking becomes steady other locomotor

patterns appear to occur in clusters: it seems that

the capacity to perform various gaits develops over

a brief period and that the occasion of the first

performance is simply subject to circumstances.

Trotting, running, galloping, and jumping were

observed for the first time in all animals between

the ages of 20-30 days, the unsupported bipedal

posture between 30-40 days, and vertical looping

(see "Climbing" below) up and down steeply inclined surfaces from 32-40 days. As Ducker (1965)

noted, the first climbing experiences are characterized by an unwillingness to descend without assistance. The types of locomotion in G. tigrina can be

divided into walking, trotting, running, galloping,

climbing, jumping, and postures associated primarily with all of the preceding. Animals displayed

a marked following response to the writer when introduced to unfamiliar rooms from the ages of 35-38

days (N = 2); it was impossible to evoke the re-
NUMBER 239 13

sponse thereafter, possibly because the surroundings

were familiar.

WALKING (Figure 3A).—Although walking velocity varies greatly, the footfall sequence during a

walking stride remains consistent: hindfoot, then

forefoot of one side, followed by the same order

on the opposite side. The durations (based on

8 mm film, 18 fps) of three consecutive walking

strides in an animal (O) moving moderately fast

were 0.45, 0.45, and 0.54 sec. The relatively slow

strides of another animal (K) as it rubbed its back

against the wall of the cage were 1.8, 3.3, and 2.0

sec. When it finished and walked away, four strides

measured 0.77, 0.83, 0.66, and 0.72 sec in duration.

The mechanical characteristics of the phases of

limb elevation and contact are the same in G.

tigrina as in the mongooses described by Taylor

(1970).

TROTTING.—One brief film sequence of this gait

reveals a resemblance to that described for the

white-tailed mongoose (Taylor, 1970); the footfall

sequence is left hind, right forefoot followed by

right hind, left fore; however, the hindfoot-forefoot

sequence is not inflexible according to Muybridge

(1957). In my film there is no contact-free suspension phase. At times, though, the trot appears to

be sufficiently fast that two support-free transit

phases do occur per stride. The duration of one

trotting stride in O was 0.23 sec.

RUNNING.—Running and rapid trotting are identical, as both involve diagonal limb extension with

two support-free transit phases. Though no film

adequate for an analysis of running was made, it

appears that as a gait it is secondary in importance

to galloping and may occur most frequently as a

transitional gait between walking or slow trotting

and galloping.

GALLOPING (Figure 3B).—In this gait, thrust is

provided alternately by the forelimbs and hindlimbs with the latter supplying the greatest propulsion. The interval between forelimb elevation and

footfall of the hindlimbs is extremely brief. During any one galloping sequence, one foreleg usually

predominates as the leading limb; but occasionally

apparent simultaneous footfalls occur, as do transitions to an opposite leading limb. Of 77 galloping strides (from 11 incompletely recorded bouts),

60 strides were led by the left forelimb, 12 by the

right, and in 5 both forelimbs appeared to strike

the ground simultaneously.

CLIMBING.—Genets are adroit climbers able to

traverse horizontal surfaces in an inverted position

and to ascend and descend surfaces inclined up to

and greater than 90° from the horizontal. All gaits

can probably be used in ascending and descending

gently inclined planes; however, only two gaits are

employed in climbing up and down flat or contoured surfaces inclined at greater than 45° with

the horizontal.

A walking gait can be efficiently used on any

vertical surface that permits a secure purchase with

the claws or toes. Vertical and steeply inclined wire

mesh as well as suspended lengths of burlap are

easily ascended in this way. Walking descent however requires an irregular surface on which the

pads of the manus can partially support the hanging weight of the body. Consequently, in descending smoothly contoured objects and burlap sacks,

an alternating antero-posterior limb extension is

employed, termed "vertical looping" by Taylor

(1970). By arching the spinal column and alternately sliding the fore- and hindfeet forward the

animal is able to descend while maintaining quadrupedal contact with the substrate during most of

each stride. The same method can be used in

climbing upwards with or without the assistance of

the claws. The limits of the genet's climbing skill

are tested when ascending a smooth and narrow vertical surface, such as the edge of a house door. In

this situation the majority of the body weight is supFIGURE 3.—Two gaits of the genet: A, walking in a year-old animal (animal O; 24 fps—the head

is depressed in frames 8-12 as the animal smelled the substrate); B, galloping in the same animal at the age of 10 weeks (64 fps).
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ported by the forelimbs; these are adducted with

the forefeet opposed and exerting a compressing

force. The hind feet prop the hindbody out so

that the force vector acting on the forelimbs is at

a tangent with the vertical. As the hindlegs extend,

the forefeet release their grip and rapidly seize an

elevated purchase. Flexion of the spine and hindlegs then permit the hindfeet to assume progressively higher grasps. Vertical looping under these

circumstances is considerably strained, for, unlike

a branch, the smooth parallel surfaces of a door

do not permit the use of the claws or an embracing

type of purchase. Because climbing in an inverted

horizontal position requires three feet in contact

at all times, only the walking gait can be used.

JUMPING.—This movement consists of four successive phases and usually involves a limb coordination similar to that of the gallop. The jump is

often preceded by a quadrupedal stationary phase

characterized by retracted mystacial vibrissae, visual

fixation of the landing target, and intention movements expressed by alternating up and down motions of the forebody and adjustment of foot position. The length of the metatarsus touches the

ground and the hindlegs are strongly flexed (Figure

4A). The propulsion phase begins with a push by

the forelegs that apparently guides the torso into

a position where the hindlegs can be straightened

with maximum effectiveness (Figure 4B). The

vibrissae are extended foreward, and, as the hindlegs propel the body up and forward, the forelimbs

swing into an extended position (Figure 4c). During

suspension-free transit, the third phase, the hindlegs

partially flex beneath the body. On landing the

forefeet strike first, but unlike galloping they usually hit simultaneously (Figure 4D). Landing impact forces the forebody down so that the vibrissae

often contact the substrate. Contact of the hindlegs

is preceded by a strong flexion of the upper forelimb segments, a forward shift of the torso, and a

flexing of the spine (Figure 4E). The entire metatarsus makes contact during landing, and momentum forces the body forward several steps.

In jumping both long horizontal and vertical

distances the forelegs usually land on the top of the

object, but the hindlegs often strike against the

side of the object and then with a walking motion

propel the hindquarters up. In short jumps, none of

the limbs may land simultaneously (Taylor, 1970).

Occasionally and especially during intense social

interaction, genets jump upward from a bipedal

stance and land on the fore- and then the hindfeet,

or with a variable footfall sequence. They can

also jump backwards.

Hall-Craggs (1965) states that Galago senegalensis

maintains a gaze fixed on the landing target at

least until leaving the ground. During the photographing of Figure 4, the electronic flash was once

positioned in front of the jump path and discharged several times when the genet was in midair.

The animal often over- and undershot its landings,

suggesting visual feedback is important during the

jump.

GAIT-ASSOCIATED POSTURES.—Though gaits can be

described in terms of the phase relationships of the

limbs and the duration of the stride and its components, posture is often overlooked unless it is an

inherent property of the gait. The ability to maintain a more or less constant spinal contour is

inversely related to the speed of the gait. While fast

gaits involve rapid, extensive, and recurrent changes

in the spatial configuration of the vertebral and

limb segments, slow gaits confer few constraints on

the spinal column, and the phase relations of the

limbs are relatively unaltered over a broad range of

limb flexion and extension. A graded continuum

of walking postures exists from a low crouched

posture to a strong arched-back posture (Figure

5B-E).

Based on still photographs and film, Figure 5

depicts four variants in walking posture and the

immobile extremes at either end of the series. In

crouched walking the head and neck are depressed,

the humerus and femur strongly flexed, and the

elbow and knee act as the primary moment arms

(Figure 5B). During moderate flexure of the back

there is minimal bending of the leg joints, and the

heads of the humerus and femur appear to be the

main moment arms (Figure 5E). Though a few

short steps may be taken in the extreme postures

(Figure 5A,F) slow sustained locomotion does not

appear to be possible unless the stances are modified

to approximate the next most similar postures.

In addition to the postures associated with gaits

that often occur in a social context, a crouched

elongated posture is assumed during an irregular

walking approach toward novel objects. Once an

object is in range, the animal shifts the trunk forward between the limb girdles and leans toward

it. During the approach and leaning phases, the
FIGURE 4.—Phases during jumping in the genet: A, quadrupedal stationary phase

(take-off position); B, propulsion; c, suspension-free transit; D, landing (simultaneous

touchdown of the feet); and E, landing (forward swing of the torso and flexing of the

spine).
16

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

B

FIGURE 5.—Postural variation associated with stationary body

positions and the walking gait in the genet: A, retracted-Iimb

ventral reclining position; B, crouched fore- and hindquarters,

minimal spinal curvature; c, normal walking and standing;

D, semicrouched forequarters and semiraised hindquarters,

moderate spinal curvature; F, semiraised fore- and hindquarters, moderate spinal curvature; F, maximally raised

fore- and hindquarters, maximum spinal curvature, A and F

are stationary body positions; B-F can also be tri- or quadrupedal stances; B-E are also assumed during walking (drawings from photos and movie film).

FIGURE 6.—A startle or sudden change in body position in the

genet (animal £; 24 fps); within less than a half second £

made half a reversed walking stride and became motionless

in a crouched quadrupedal stance.

direction of movement often vacillates, but usually

no more than several steps are taken in the reversed

direction.

Startling produces a sudden change in body position followed by flight or immobility. This is often

succeeded by a crouched stance in which the legs

are spread and the ventrum may touch the ground

(Figure 6).

SHAKING AND TWITCHING MOVEMENTS

HEAD AND BODY SHAKING (Sichschutteln).—Like

many other patterns, shaking is a complicated motion of graded intensity. It is apparently accomplished by rapid and alternate partial rotation and

lateral bending of the vertebral column with a

resultant abrupt tossing of the overlying skin and

hair. The head and neck; head, neck, and shoulders; or the torso can be shaken independently; or

the whole body is shaken with an antero-posterior

wave of emphasis (Schutteln des ganzen Korpers).

When head shaking accompanies or precedes

shaking of the neck and shoulders, its motion is

primarily rotatory because the angle between the

head and neck is small. When only the head is
NUMBER 239 17

shaken, its movement is pendulous because the

occipital angle roughly approximates 30 or more

degrees. Shaking of the torso seems to be facilitated

by a simultaneous circular motion of the vertebral

axis that is sufficiently forceful to cause the hindlegs to take one or two steps in place. An analysis

of slow motion film would greatly help our understanding of these involved movements.

Shaking of the head and body appears to be

a response to direct tactile sensation of the skin and

perhaps indirect dermal stimulation mediated by

disturbance to the nap of the hair. The patterns

are sometimes seen after an animal has rolled and

rubbed on the ground, and are almost always seen

when the fur is saturated with water. Thus, it

serves the removal of water and particulate matter,

and may also create a uniform nap in the pelage

(Leyhausen, 1956). In addition, head shaking is

evoked by presumably painful stimuli to the face,

ears, and mouth, and may be considered as a component of the facial protective response described

by Andrew (1963). It has been observed, for example, after an animal struck a hard object with its

open mouth, received a bite on the face, or had

mystacial vibrissa pulled out by another animal.

Moreover, it is elicited by repeated mild tactile

stimulation to the ear (see "Ontogeny of Shaking,

Stretching, and Self-Directed Behaviors in Genetta"

below) and once as a bolus of regurgitated food

passed from the esophagus into the mouth.

LIMB TWITCHING.—Both the fore- and hindfeet

are twitched and shaken. The movement, often

occurring in bi- and tripedal stances or during

walking, involves a slight flexion of the limb

followed by a rapid snapping extension and retraction (Figure 7). The result is a complete but momentary straightening of the wrist or ankle and a

partial extension of the elbow and knee. If the

motion is repeated in rapid succession, the limb

has the appearance of being shaken.

Animals are often seen to twitch their hindlimbs

when their feet become wet from the puddle created

during urination. At other times, when the body

is completely wet, each foot is often raised and

shaken. Animal K shook and twitched its forefoot

repeatedly after being bitten between the toes by

a rat it was attempting to catch. In addition, the

forefeet and, to a lesser extent, the hindfeet sometimes twitch spontaneously while an animal is

asleep or resting. Forelimb twitching has also been

2.4

FIGURE 7.—Twitching of the forelimb in the genet (animal

E; 24 fps): in frames 1 and 2 the wrist is flexed; in frame 3

it is rapidly extended and in frame 4 it snaps back into a

flexed position. The twitching of this leg occurred as it

moved a step forward.

seen in a variety of social situations that can be

subjectively interpreted as containing elements of

surprise, apprehension, or conflict.

STATIONARY BODY POSITIONS

Stationary body positions may occur as relatively

short-termed interruptions of locomotion, or more

frequently as protracted periods of rest. Although

"posture" is the term generally used for motionless

body attitudes, it will be used herein to refer to

configurations of the spine and limbs, and positions

of the soft anatomy and hair at any time irrespective

of locomotion. This is consistent with its general

meaning of "relative arrangement of the different

parts," and "state and condition at a given time."

 s

In the following discussion the term "stance" will

be used for the basic types of foot position assumed

by immobile standing animals. The term "reclining

position" is self-explanatory. The descriptions under

these categories are not equally complete because

the attitude of the entire body is considered in

reclining position, while only foot position is necessary to define a stance. Suffice it to say that a range

• Webster's Seventh New Collegiate Dictionary, 1963.
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

of spinal curvatures is possible during quadrupedal and tripedal stances. The list of reclining

positions includes those most commonly seen and

does not consider postures that result from spatially

confining circumstances.

QUADRUPEDAL STANCES.—The animal stands with

the four legs providing body support. In an upright

quadrupedal stance the forelegs rest against an

elevated surface, the trunk of the body is inclined

upwards, and the weight is mainly sustained by the

hind feet.

TRIPEDAL STANCES (indecision-alert posture:

Ewer, 1968).—When the body is supported by three

legs, it is usually one of the forelegs that is partially

flexed and elevated. A hindleg is rarely elevated for

any length of time. The stance occurs most frequently during pauses in slow walking, and essentially constitutes an arrested walking stride.

BIPEDAL STANCES (Mannchenmachen).—Three

variations of bipedal body support have been observed. Support may be provided by the toes and

interdigital pads (high-sit: Ewer, 1963), by the

entire metatarsus (semibipedal: Ewer, 1968), or by

the metatarsus, the rump, and the tail base (low-sit:

Ewer, 1963; Figure 8A,B). All variations can be

maintained for up to several minutes.

SITTING.—This posture is characterized by three

zones of contact with the substrate: the length of

the tail, the length of the metatarsus, and the forefeet; the contact zones can be maintained with

considerable variation in forelimb flexion (Figure

8c). Occasionally one foreleg may be elevated.

VENTRAL RETRACTED-LIMB RECLINING POSITION

(aufrechten liegen mit Vorderpfoten nach innen:

Leyhausen, 1956).—In addition to the three zones

of contact that characterize sitting, the chest and

belly and often the radio-ulna also touch the

ground. The head may be elevated or may rest on

the forelegs or the substrate beside them (Figure

8D).

SEMILATERAL, SEMIEXTENDED RECLINING POSITION.—This position differs from the retractedlimb reclining position in that the hindlegs are

usually extended and the lateral surface of one leg

contacts the ground broadly (Figure 8E).

EXTENDED BODY RECLINING POSITION (ausgestreckte Ruhelage: Goethe, 1964).—In these positions, the legs are more or less extended, at least

not flexed against the body. Usually the animal

reclines with either the ventrum or one side of the

body contacting the substrate (Figure 8F). Dorsal

extended body reclining is less frequently seen and

cannot be maintained without lateral support.

LATERAL CURLED RECLINING POSITION.—The torso

is flexed, limbs are usually partially flexed, and one

side of the body broadly contacts the ground. The

head usually rests on or between the hind legs and

is covered by the tail which is wrapped about the

body contour.

ASSUMPTION OF RECLINING POSITIONS.—Unlike

ungulates viverrids have low centers of gravity and

versatile limb mechanics that impose few restrictions on the method of reclining or standing

(Zannier-Tanner, 1965). Retracted-limb reclining,

for example, can be assumed by simultaneous

lowering of fore- and hindquarters, or by lowering

the forequarters from a sitting position. Sitting is

often the transition attitude between reclining and

standing. Circling in place, characteristic of canids,

often precedes the assumption of reclining postures

in which the hip and thigh contact the substrate.

In these positions the animal first depresses the

rearend and then sits on the side of the hip.

STRETCHING MOVEMENTS

WALK STRETCH (Schreitstrecken: Eibl-Eibesfeldt,

1951).—This protracted and exaggerated form

of walking varies considerably in both duration

and complexity. In its most complete form it involves three phases: (1) a usually unequal anterior

extension of the forelegs, extension of the neck,

and a slight concave arching of the back; (2) an

extension of the hindlegs, depression of the hindquarters, and an accompanying shift forward of the

pectoral girdle; loss of the spinal concavity and the

onset of a ventral tailbase flexion; and (3) continued

tail flexion as the feet make several short stiffjointed steps. Partial versions of this stretching

movement omit either the first or the latter two

phases. When initiated from a sitting position, the

stretch may involve only the first phase but without

the spinal concavity, or may continue in a standing

position to completion (Figure 9).

UPRIGHT STRETCH.—The animal assumes a quadrupedal upright position, extends the forelegs up

wards, and stretches the head and neck up and

back. The claws are sometimes unsheathed during

this movement.

ARCHED-BACK STRETCH (Figure 8c; Streckbuckel:
FIGURE 8.—Stationary body positions and stretching movements in the genet and the African

palm civet (Nandinia): A, bipedal stance with the metatarsus touching the substrate; B, with

only the toes touching the substrate; c, sitting position; D, ventral retracted-limb reclining

position; E, semi lateral semiextended reclining position; F, extended lateral reclining position;

c, arched-back stretching; H, sitting stretch; i, yawning.
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

21

30

36

FIGURE 9.—A walk stretch in the genet initiated from a sitting

position on a log (animal C; 18 fps): frames 1-7, stretching

of the forequarters (notice unequal extension of the forelimbs); frames 21-30, stretching of the hindquarters (typically

the hindlegs are unequally extended); frame 36, a hindleg is

moved forward (the tail is obscured by the animal's body).

Leyhausen, 1956).—This movement, performed in

a motionless standing position, involves extension

of the fore- and hindlimbs, a pronounced flexion of

the back, and often a dorsal curling of the tail. It is

possible that arched-back rubbing (see "ObjectOriented Contact Patterns" below) is also a modified type of arched-back stretch.

SITTING STRETCH (Figure 8H).—The forelimbs are

extended and the scapulae elevated while the neck

is ventrally flexed and the head pointed down and

the chin tucked in.

TAIL STRETCH.—The base of the tail flexes ventrally while the distal one-half to two-thirds extends

posteriorly. Tail stretching is usually performed

concurrently with the walk stretch.

YAWN (Gahnen).—In its most complete form

the jaws are gaped maximally to about 80 degrees,

the head tilted upward, the vibrissae extended, the

eyes closed, and the tip of the tongue curled. In

its weakest expression, the jaw gapes to about 30

degrees and the other components are absent (Figure 8i).

OBJECT-ORIENTED CONTACT PATTERNS

METATARSUS SCUFFING (Figure 10).—In a posture

similar to sitting (with the metatarsus but not the

tailbase contacting the substrate), the hindfeet alternately stroke backwards with the pads scuffing the

surface. The claws are not extended, and the feet

may or may not be lifted when brought forward

FIGURE 10.—Metatarsus scuffing in the genet (animal K; 24

fps): frames 1-6, the visible leg is thrust posteriorly and the

metatarsus dragged on the substrate; frame 9, the leg returns

to its initial position.
NUMBER 239 21

again to their original position. During this movement the body may be stationary or may be slowly

propelled forward by small steps. An animal was

once seen to scuff with one hindfoot elevated on a

horizontal log. Scuffing was most frequently seen

performed within and on top of plywood and cardboard boxes. Occasionally a rough-surfaced log was

used.

SUBSTRATE-CLAWING (Krallenscharfen: Leyhausen, 1956).—The claws are extended and hooked

into the substrate as the legs are tensed and slightly

retracted in an often sporadic, but alternating

rhythm. Clawing is usually performed in a crouched

posture on horizontal and diagonal branches, and

involves either the forelimbs or both pairs of limbs.

Often the claws are not dragged across the surface,

but appear to be tensed as the limbs are flexed.

As differential growth of the dorsal and ventral

surface of the claw account for its recurved and

pointed structure, it seems unlikely that substrateclawing actually sharpens the claw. Being retracted

most of the time, it furthermore experiences relatively little wear. Rather, it appears that substrate

clawing removes loose fragments of keratin and claw

sheath, and also keeps the claw worn. It is well

known to zoo keepers that a civet or cat that lacks

a suitable clawing substrate will develop long claws

which can penetrate the toe pads and become

ingrown.

ARCHED-BACK RUBBING (Figure 11).—Though this

behavior resembles in posture the arched-back

stretch, it differs in that the animal leans against a

vertical or diagonal surface while slowly walking

forward. The movement typically begins with a

strong central flexion of the back. As the animal

steps forward the posterior half of the trunk attenuates, and the flexion shifts anteriorly. When the

hindquarters are brought forward again the arch

assumes a more central position. The areas rubbed

are the hip, side, shoulder, neck, and occasionally

the side of the head.

HEAD AND NECK RUBBING (Figure I 5B).—The

variety of stationary body positions assumed during

head and neck rubbing depend upon the location

of the object being rubbed. If the animal effects

this on a horizontal or diagonal surface, a sitting

or flexed-limb reclining position is adopted; upright

objects are rubbed in a quadrupedal upright position. Rubbing of the head and neck is achieved by

a combination of rotating, extending, and flexing

FIGURE 11.—Arched-back rubbing in the genet (animal K; 18

fps): frames 1-21, changes in the extent and center of spinal

curvature; frame 21, the animal is beginning to lick its breast.

movements directed usually to the substrate and to

targets on the animal's own body (forelegs and tail).

Though the sequence and form of the movements

are highly variable, a fairly distinct component of

head and neck rubbing is a strokelike head extension in which the lips on one side of the rostrum

maintain contact with the substrate.

Certain aromatic substances are especially effective in eliciting this behavior. Animal O would

consistently rub his head and neck against my hair

when it was scented with hair oil. On these occasions the animal maintained a supported quadrupedal stance on my shoulders, clasped my head

with its forepaws, and alternated the rubbing with

Flehmen and licking my hair. (See "Sniffing" under

"Companion-oriented Contact Patterns" below.)

RECLINING, RUBBING, AND ROLLING.—Though
22

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

genets occasionally lie down and rub and roll with

sinuous movements of the spine similar to a dog,

the behavior is performed most frequently if the

animal's fur is saturated with water. In addition, a

wet animal often propels its body forward with the

hindlegs while sliding the side of the head, neck,

and shoulder on the substrate.

PERINEAL-GLAND MARKING.—All of the postures

employed during the apposition of the perineal

scent gland to the substrate are characterized by

strong flexion of the hindlimbs. When the gland

is pressed against a low-lying object, the squatting

position may be quadrupedal or tripedal. In fourlegged squatting the hindlegs are level or one is

placed on an elevated object; in three-legged squatting the elevated leg is flexed. In marking elevated

vertical objects or the undersides of elevated

horizontal and diagonal surfaces a reversed upright

quadrupedal position is used. In all positions the

appression of the perineal gland to the object may

be accompanied by a rhythmic lateral pelvic oscillation. Though both sexes presumably have the

ability to mark by all methods, only males have

been seen to use the "handstand" position; and,

with one exception, only females have been seen

to mark in a squat position. The presence of elevated scent marks in cages occupied only by females,

however, suggests the handstand position is not

sex specific. The gland is everted shortly before or

as it is apposed to the object. Eversion also occurs

when animals are restrained by hand or held by the

tail and, at least in females, eversion accompanied

by a lordotic posture can be evoked by rubbing the

area between the labia of the relaxed gland.

The gland is a longitudinally folded muscular

pad having the appearance of two densely-furred

labia situated between the anus and vulva or the

scrotum and penile aperture. With muscular contraction of the gland the labia part and the inner

surface everts to form a firm, raised, oval-shaped

pad covered with a fine layer of white hair (Figure

12). The median sulcus, a shallow longitudinal depression corresponding to the floor of the folded

gland, is bordered anteriorly by a deeper Y-shaped

crease, and centrally and posteriorly by transverse

creases. According to Pocock each crease on either

side of the median sulcus forms a crypt into which

scent, produced by underlying glandular tissue, is

secreted via "a pair of laterally placed clusters of

minute orifices" (1915c: 190). Some species apparFIGURE 12.—The perineal scent-gland of the genet: A, closed

condition; B, everted condition, top row showing profiles of

the closed and everted gland (po = penile opening, gf =

glandular fold, t = testes, a = anus, ms = median sulcus.

Is = lateral sulcus).

ently exhibit sexual differences in the number of

secretory crypts (Pocock, 1915c).

A clear, oily emulsion covers the inner surface of

the relaxed and folded pad, but the exudate within

the crypts often has a denser pomade-like consistency and a yellowish color. Although the entire

pad may secrete scent, it seems likely that incidental

motion of the folded labia and capillary action

distribute the contents of the crypts within the

folded pad. With repeated deposition on wood or

metal the clear exudate becomes a dark brown

coagulum. Kingston (1965) also reported this color

change for the perineal secretion of the African

civet (Civettictis civetta). Repeatedly deposited scent

can penetrate wood to a depth of several centimeters (Figure 15c) and retains its redolence for at

least four years. The odor is a mildly sweet musk

that according to D. Miiller-Schwarze (pers. comm.)

resembles civetone from which impurities such as

skatole have been removed.

On three occasions when females displayed signs

of sexual receptivity, both an increased frequency

of scent-marking was observed, and the rooms in

which they were caged manifested a heightened
NUMBER 239 23

scent of urine. Over a 19-day period in which 90-

minute observations were made on alternate evenings, one female (C) exhibited an erratic pattern

of intense marking (Figure 13). Curiously, this

pattern does not seem to reflect a relationship to

the gradual hormone changes characterizing the

estrous cycle. Furthermore, the peak in marking

occurred after the apparent height of estrus. More

significant perhaps is the observation that the majority of scent-marking did not occur in the immediate context of scent-sniffing. This suggests that in

this situation the odor of scent perceived at close

range is not important in releasing or orienting the

marking act. A heightened rate of secretion during

estrus probably also contributes to the heavy scent

accumulation at marking sites. These sites become

encrusted with urine and it is my impression that

both urine and vaginal secretions are deposited as

the perineal gland is rubbed. A drainage of fluid

from the vagina into the perineal fold would seem

to be a natural consequence of the contiguity of

these two organs. Estrous females also dribbled

urine on scent marks independent of rubbing the

perineal gland.

ANAL DRAG.—The animal assumes a sitting posture with partially spread hindlegs and walks forward with small steps dragging its anus on the

9 11 13 15 17

DAYS

FIGURE 13.—Frequency of perineal scent marking and scent

sniffing in a female genet (animal C) on days before, during,

and after sexual receptivity (shaded bar — scent marking;

white bar = scent-mark sniffing independent of scent marking; black bar = scent-mark sniffing following scent marking;

a = female observed in copulo, presumably indicating day of

peak estrus; b = female observed mounting her cagemate).

ground. Though dragging of the perineum in a

similar manner has not been observed, genets seem

to be physically capable of modifying the emphasis

of the movement so that this is possible.

FEET.—The forefeet are used in several ways to

contact and manipulate inanimate objects depending on the object's situation. An animal may

touch an object while standing, sitting, or reclining

on its side or back by extending a foreleg and

contacting the object with the pads of the pronated

or semisupinated forepaw. The object may be

drawn toward the body by hooking the paw slightly

and sometimes extending the claws while retracting

the limb. A combination of adduction and retraction of the extended limb may result in weakly

pushing an object aside. For instance, I have seen

an animal push a large cardboard box away from

a wall. For this the limb always moves the object

toward the opposite side of the body. Though

usually seen as a companion-oriented contact pattern, clasping with the fore- and hindfeet occasionally involves inanimate objects. When the

forelegs are used to clasp, the paws are almost

always semisupinated. For clasping an object the

attitude of the hindlegs is the same used for

climbing down a narrow vertical log; the pads are

completely or semisupinated with the object held

between or pressed against the belly.

NOSE.—Often the rhinarium lightly touches the

surface an animal is smelling, and leaves behind

a trace of watery mucous. The ventro-anterior

portion of the rhinarium together with the upper

lip on either side of the philtrum and possibly the

tip of the lower jaw form a zone that is quite sensitive to tactile stimuli. This area is used to lightly

touch objects being smelled, or to playfully push a

small object along the floor. On a few occasions an

animal was seen forcefully pushing its rhinarium

against a novel object; such behavior is far more

prevalent in Civettictis than in Genetta.

OBJECT-ORIENTED CONTACT PATTERNS IN OTHER

GENERA

PERINEAL-GLAND MARKING.—The scent-marking

gland in Civettictis displays a number of distinct

differences from that of Genetta. It is proportionately larger and occupies a slightly more posterior

position; like the gland of Viverra zibetha described

by Pocock (1915c), the lips are confluent, partially
24

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

fused anteriorly, and divergent posteriorly. The

gland's pouchlike character prevents its complete

evacuation during scent marking, and consequently

some of the contained civetone becomes dark prior

to marking. The strong, mildly fetid odor of the

scent disappeais after several days of exposure.

Wood shavings from the cage floor occasionally

became lodged within the female's pouch, where

they remained for at least several hours before

being deposited with a scent mark.

A normal quadrupedal stance is employed by

male and female Civettictis to mark surfaces at

the level of the perineal gland. The gland is usually

everted shortly before or as the rear end is turned

toward the marking object; one or two steps are

taken backwards; the tail is elevated and/or

deflected laterally, and the gland is pressed against

the surface for two seconds or less, usually without

accompanying pelvic motion. The female, however,

was once seen executing a weak lateral oscillation

against a 1 cm mesh wire, and sometimes dipping

the rear end slightly as she ended glandular contact.

The gland usually closes as soon as contact terminates. Though Civettictis have not been seen marking low-lying objects in a squatting stance, scent

marks on a cement floor (Figure 15H) may have

been deposited in this way. They may also have

been made passively as the animal was sitting.

Nandinia and Paradoxurus have been observed

dragging and wiping (without locomotion) their

scent glands on inclined and horizontal logs. While

these methods of marking differ from one another

in duration, both show only minor differences in

posture from dragging and wiping the anus. Unlike

Genetta and Civettictis, the glands in these genera

are thin walled, hairless, and devoid of secretory

accumulation. Nandinia's pregenitally situated

gland can be freely opened into a flat, elongate

oval area even as the animal climbs about. This

sporadic opening and closing is not associated with

scent-marking but may be a passive response to

incidental skin movement. On a few occasions when

the male became excited a sweet odor was detected,

but it is unknown whether or not the gland was

the source. In Paradoxurus and Paguma the gland

has a circumgenital position with the penis and

vulva opening centrally and the labia folding

posteriorly.

The male and female Hemigalus and the three

female Galidia were observed occasionally wiping

the hindquarters both spontaneously and after urination, but whether these movements resulted in

the deposition of specialized dermal gland exudates

is unclear. Gregory and Hellman (1939) briefly mention the existence of small perineal glands within

the Hemigalinae, but more precise information on

their occurrence and structure apparently remains

to be discovered. Likewise, little is known concerning the perineal glands of the Galidiinae.

NECK-SLIDING (Figure 14).—Civettictis, Nandinia,

and Paguma exhibit this behavior with only minor

differences. It is always preceded by sniffing the

object or area to which the body contact is directed.

Typically the animal stands with the head over

the object and simultaneously depresses the forebody, partially rotates and turns the head, and

slides the posterior cheek region or neck over the

target until the shoulder approaches or touches it.

Foreleg support of the body is sometimes relinquished and the animal slides onto the shoulder.

The neck and forebody is then raised, the head

repositioned over the target (which may again be

sniffed briefly), and the motion is repeated though

frequently on the opposite side of the neck.

The main differences between the neck-sliding

pattern in Nandinia and Paguma, on the one hand,

and Civettictis, on the other, involve the stance

and the extent of the forebody that contacts the

target. While the palm civets often depress the

hindquarters during neck-sliding, the rump of

Civettictis is held almost as high as in normal

standing. In consequence of its higher center of

gravity, Civettictis sometimes tends to lose its

balance once it slides onto its shoulder. Rather than

falling over, however, the animal usually quickly

regains a quadrupedal stance. Sliding onto the

shoulder is infrequent in the two palm civets, and

when it does occur contortion of the torso is less

exaggerated. In addition, Civettictis occasionally

initiates neck-sliding by anointing the length of the

throat with a few anteriorly directed and unrotated

extensions of the neck. Thus the sides of the neck,

the shoulder region, and to a lesser extent the

throat are the chief anointing targets in Civettictis,

while the posterior cheek region and the sides of

neck are primarily marked in Nandinia and

Paguma.

Neck-sliding seems to be both released by and

oriented to certain odors, but the source need not

be tactily or visually discernible. The high degree
FIGURE 14.—Neck-sliding in the African civet (Civettictis): A, B, frontal views of alternate sides

of neck-sliding over a sponge soaked in a lemon-scented detergent; c, sliding onto one shoulder

(S feet and the neck-shoulder are supporting the body); D, sliding without foreleg support.

of target specificity results from the initial sniffing

orientation, and also, but not necessarily, from

contact with the object. Both Nandinia and Paguma

executed neck-sliding upon dead rats and mice and

in their own and other animals' (civet, genet) urine

and feces. Diicker's (1971) male and female Nandinia would slide the chin, neck, and sides of the

head, as well as the breast for up to 15 minutes on

a board that had been used for cutting liver. This

behavior was also elicited by solutions of heliotrope

(Baldriantinktur) and cologne.

On several occasions after the male Paguma

killed large (400 g) rats it directed its neck-sliding

to the prey's lower abdomen and genitalia which

forced a flow of urine from the carcass. In these

instances the animal's activity was clearly released

by and oriented to the smell of the anogenital

region; the crushed heads of the prey were left

alone, though they certainly provided a distinct

alternative scent. Dead smelt, mice and rats, live

mouse pups, genet urine, and a lemon-scented dish

detergent were all highly effective elicitors of this

behavior in Civettictis; occasionally a piece of

horsemeat worked. All species sometimes lick the
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

neck-sliding target, and if the target is food it is

usually ingested afterwards.

The effect of the neck motion on the scent

source varies. Soft substances and liquids impregnate the hair as they are spread upon the substrate.

The bodies of mice, rats, and fish sometimes remain

stationary as the neck slides over them, but more

often they slide or roll especially if their axis is

parallel to the direction of the neck movement.

The pressure exerted by the neck upon the scent

target must be considerable. Smelt are sometimes

broken into several pieces, and mouse carcasses are

noticeably distorted. Mouse pups are crushed by

neck-sliding and their blood stains Civettictis'

white neck stripes. After an animal neck-slides on

animal carcasses, they are usually eaten.

UPRIGHT QUADRUPEDAL VENTRUM-RUBBING (Abreiben in hochaufgerichteter Stellung: Vosseler,

1929).—Cryptoprocta assumes an upright quadrupedal stance in order to rub the ventrum (from the

sternum to the pelvis) on vertical surfaces. The

sides of objects, such as vertical tree trunks, are

grasped with the forefeet while the hind feet rest

on the floor on either side of the trunk, alternately

extending and flexing to produce an up and down

rubbing motion. The inguinal region received the

majority of the rubbing in the male animal

observed, while Vosseler (1929) saw two males and

a female rub the breast to the inguinal region on

cage projections such as metal bars, and the edges

of planks and walls. An abundant and easily removable fatty secretion on the male's rust-red

colored ventrum fur suggested to Vosseler the existence of a diffuse dermal scent gland, but no macroscopic or histological investigations have been made.

In the male Cryptoprocta I observed, ventrumrubbing was almost always preceded by several

sequential or simultaneous activities. Most commonly the tree trunk was clawed with the forefeet

and the cheeks and lips on alternate sides of the

head were rubbed back and forth. At times headrubbing was interspersed with chewing on the trunk

and branches. Under some circumstances, as when

a new branch was placed in the cage, chewing

became intense (J. F. Eisenberg, pers. comm.). Also,

urine was sometimes dribbled during these activities, and on one occasion clawing, face-rubbing, and

urination occurred simultaneously. The upright

quadrupedal stance assumed during these activities

differed from the position used during ventrum

rubbing in that the hindfeet were placed several

decimeters away from the trunk, and the back was

bowed.

SIDE-RUBBING.—The female Nandinia was often

seen rubbing one side of its back up and down

by leaning against a screen wall in the cage and

extending and flexing the legs, particularly the

hindlegs. Besides massaging the skin, the movement

also removed loose hair that accumulated at each

rubbing site.

COMMUNICATIVE FUNCTION OF OBJECT-ORIENTED

CONTACT PATTERNS IN Genetta

The tendency for animal companions to smell the

often specific sites where body contacts are repeatedly oriented suggests that chemical communication may be an incipient or definitive function of

certain object-oriented behaviors (Eisenberg, 1963,

1967). Consideration of the spatial and chemical properties of the odorous substances deposited

during these activities provides a convenient though

subjective method of evaluating the communicative potential. Table 4 outlines and ranks some of

the features of deposits that result from objectoriented patterns.

The residue of the perineal gland (presumably

civetone) is subjectively the most potent source of

chemical information, for its redolence persists and

it is deposited in combination with other substances. At least in caged animals, scent deposits

display a bimodal and sometimes discontinuous

vertical distribution. This is undoubtedly affected

by cage structure and the scarcity of suitable

objects, but the difference is also a partial function

of the two basic stances assumed during marking.

In the reversed upright quadrupedal stance, scent

is usually deposited at an elevated position on vertical and diagonal planks, and on the undersides

of raised diagonal and horizontal logs. The mean

central heights of scent marks in cages at the

National Zoo, for example, were 22.5 cm on vertical posts (N = ll), and 30 cm on diagonal logs

(N = 3). Marks that were presumably made in a

quadrupedal squat averaged 21.2 cm on 45° plywood edges (N=9) (Figure 15D), or were limited to

a height of 4.5 cm on the baseboards of the cage

(N = 6). The length of the scent mark is likewise a

function of the angle of the object, though marks

are also enlarged by absorption into the wood. The
NUMBER 239 27

TABLE 4.—Distribution of scent marks within genet and civet cages

at the National Zoological Park (measurements in cm)

Scentmark

object

Cage

occupants

Genetta

Vertical and

diagonal

surfaces

Subtotal

Undersides

of logs

Subtotal

Total

I,

E,

c,

I ,

c,

Civettictis

Vertical

surfaces

Total

9

<S

F

0

K

F

K

Number of

scent-marks

per cage

12

6

4

22

1

2

3

25

14

1

15

Vertical distance

from substrate

Smallest

Mean

16.7

20.3

17.9

17.5

17.5

25.5

22.8

17.8

28.0

30.5

28.2

Range

(12.5-23.0)

(15.0-23.0)

( 8.0-24.5)

( 8.0-24.5)

(13.0-38.0)

(13.0-38.0)

( 8.0-38.0)

(21.5-30.5)

(21.5-30.5)

Greatest

Mean

25.9

33.8

27.1

29.6

25.5

43.0

37.2

28.3

31.9

33.5

31.7

Range

(19.0-32.5)

(23.0-44.0)

(26.0-28.5)

(19.0-44.0)

(30.5-55.5)

(25.5-55.5)

(19.0-55.5)

(30.0-34.0)

(30.0-34.0)

i o

scent-marks

Mean

9.2

13.4

9.2

11.3

8.0

17.5

14.3

11.3

3.9

3.5

4.8

Range

( 3.5-15.5)

(12.5-22.5)

( 3.5-18.0)

( 3.5-22.5)

( 8.0-17.5)

( 3.5-22.5)

( 2.5- 8.5)

( 2.5- 8.5)

longest marks occurred on horizontal baseboards

(x = 23.8 cm), medium length marks on undersides

of diagonal logs (x = 14.3 cm) and on 45° plywood

edges (x = 11.8 cm), and the shortest marks were on

vertical posts (x = 9.9 cm). By using the quadrupedal squat position an animal could potentially

mark the upper surfaces of all the elevated branches

in the cage. This, however, is not done because

scent-marking is a strictly terrestrial activity.

There obviously has been a strong selection for

elevated scent marks as a reversed upright quadrupedal stance has evolved independently in members of three carnivore lineages. In the absence of

detailed information on carnivore pheromone

chemistry and sensory physiology, two alternative

speculations can be advanced regarding function.

On the basis of Bossert and Wilson's (1963) findings,

it seems possible that under similar atmospheric

conditions the active space emanating from an

elevated mark would encompass a greater volume of

air than that from a lower mark. Likewise, the

diffusion properties and receiver's sensitivity may

be such that elevated marks increase the probability of detection by sympatric species. This is

suggested by the following observations: (1) the

level of Civettictis' marks coincides and is broadly

overlapped by the elevated marks of Genetta tigrina

(Table 4); (2) close-range sniffing of such marks

requires an upright quadrupedal or bipedal stance

in Genetta, but is achieved in the typical standing

position of the civet; and (3) Genetta can assume

a bipedal stance and thereby more than double its

detection height when scenting the air, while Civettictis is only able to increase its height by leaning

against an object in an upright quadrupedal stance.

The latter point may seem superfluous, but

Madison and Schoop (1970) recently presented suggestive evidence that displaced terrestrial salamanders facilitate detection of home-area odors by

climbing up into vegetation. If the scent of genets

and African civets carries similar territorial messages, the overlapping heights of scent marks may

be viewed as a form of character convergence

promoting interspecific spacing (Cody, 1969).

The deposition of perineal scent in combination

with urine and vaginal fluid is probably relevant

to the "exalting" effect for which civetone has been

commercially exploited. It is doubtful that the
28

chemical capacity to fix and enhance odorous components is an incidental property of civetone or for

that matter other animal musks. In nearly all genera

in which musks have been identified (Viverra,

Civettictis, Moschus, Castor, Ondatra), the glandular source is proximal to the genitalia and anus,

thereby increasing the chance for occasional mixing

of exudate and metabolites. In this connection, the

relation of the gland's proximity to the genitalia

and the nature of the sensation generated during

scent-marking merits attention. In nearly all viverrine and paradoxurine carnivores (Nandinia and

female Arctogalidia excepted), the vagina and penis

lie directly beneath the floor of the perineal fold,

and as marking undoubtedly compresses these structures, the stimulus may well have a sexual manifestation during parts of the year. In estrous females the

compulsive and spontaneous character of scentmarking appears to be a response more to fluid

accumulation within the fold and perhaps to heightened genital sensitivity than to specific olfactory

stimuli.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The characteristics of rubbing movements do not

appear to be sufficiently specialized to have welldefined chemical communicative functions. Because

rubbing removes loose hair and massages the skin

and also often produces an apparent complacency,

both grooming and comfort functions may be assumed. Nevertheless, rubbing sites are important

foci for sniffing, and the interpretation that different

patterns may produce scent traces with differing

odors is supported by (1) the finding that considerable diversity exists in the glandular composition of

mammalian skin (Montagna and Ellis, 1959), and

(2) the observation that the circumstances associated with the three rubbing movements differ.

Sniffing is associated with the performance of all the

patterns listed in Table 5, but experimental evidence is required to ascertain whether it has a

causative or directive function or is merely incidental. Probably relevant to the role of sniffing is

the action of the scent-marking movement and the

chemical natures of body and object borne substances. An exchange of odorous substances

TABLE 5.—Some characteristics of scent deposits resulting from object-oriented contact behaviors

and from elimination in Genetta tigrina (first column: + = detectable by human nose,

— = undetectable; volatile longevity ranked in increasing order of probable magnitude)

Behavior

Metatarsus

scuffing...

Arched-back

rubbing

Head and neck

rubbing

Perineal

gland

marking

Urination

Defecation...

Anal drag

Olfactory Visible

trace deposit

-

+

+

+ +

+ +

+ +

+ +

Volatile

longevity

1

2

2

5

3

4

3

Locus Distrispeci- bution

ficity of scent

? Diffuse

+ Diffuse

Localized

+ Localized

+ Localized

+ Localized

Localized

Volatile

components

Sebum, sweat

Sebum, sweat

Sebum, saliva,

food, and mucous traces

Civetone, vaginal

secretions,

traces of

sebum, urine,

and saliva

Urine, vaginal

secretions

Feces, trace anal

scent (unknown)

Feces, trace

sebum

Associated

activities

Arched-back

rubbing

Arched-back

rubbing

Feeding, selfdirected

movements

Locomotion

olfactory

testing

Trauma, diverse

associations

Diverse

associations

Defecation,

diverse associations
NUMBER 239 29

undoubtedly occurs during any contact between

body and object. The action may be reciprocal

or may have a primarily anointing or marking effect

depending upon the chemical character of the

substances involved and the olfactory sensitivity

of the message recipient.

Head and neck rubbing, for example, accompanies three distinct activities. It occurs between

bouts of self-directed licking, as a response to

water on the fur, and to certain aromatic substances. Though the head and neck motions are

similar in all three circumstances, in the first two

instances it is oriented to both the ground or

branch upon which the animal is resting and to

the breast, forelegs, and tail. In the third situation,

Flehmen is displayed as the animal licks the scent

source, and its stance is modified to facilitate contact with the area. Thus, in the first two situations

there is probably reciprocal exchange while in the

last situation an anointing function accrues to the

rubbing.

Also, arched-back rubbing and metatarsus scuffing

sometimes resemble head and neck rubbing because

a reciprocal exchange of odorous substances occurs

and a diffuse and faintly scented field remains. The

causation of these two patterns is quite different.

Both activities occur in solitary animals either

spontaneously or as responses to tactile and/or

olfactory properties of the surroundings, and as

apparent responses to certain of the companionoriented behaviors of associated animals.

In view of the fact that all of the patterns listed

in Table 5 are regularly exhibited by solitary animals, the communicative potential of the patterns is

to an extent dependent upon the quantity of the

odorant substance deposited and its volatile longevity. Whereas urine, feces, and perineal scent are

durable, the glandular exudates laid down by the

three rubbing patterns are probably easily vitiated

by climatic conditions. It is possible that chemical

evanescence conveys recency of physical presence

and hence proximity of the marker. A consideration

of the features listed in Table 5 suggests that the

chance for a genet to detect an unfamiliar animal's

rubbing area would be rather fortuitous. It is possible, of course, that optimal rubbing targets may

be in sufficiently short supply that the use of them

may be communal. The collective contribution

would magnify the strength of the odor and visitation to rubbing loci would become traditional,

thus alleviating total dependence upon olfactory

localization.

SELF-ORIENTED CONTACT PATTERNS

MOUTH.—In genets three types of self-directed

mouth movements are commonly seen. Two of

these, licking and nibbling, often occur in alternating bouts during extended grooming sessions.

Licking directed to the trunk and limbs involves

tongue extension and retraction coordinated usually

with an upward motion of the head; the head bobs

down and up with the respective extrusion and

retraction phases of the tongue. In nose licking

(Nasenlecken: Leyhausen, 1956), the tongue slides

out, up, and over the rhinarium and is retracted

over the same path. In licking the rostrum (lips or

"chops"; Lippenlecken: Leyhausen, 1956), the

tongue extrudes anteriorly (often above the canine)

and glides posteriorly to wipe the mystacial vibrissae and the upper lip musculature. The length of

lip that is wiped varies, but the direction seems

always to be posterior and may extend as far as the

maxillary spots in front of the eye (Figure 15F).

Nibbling (Beknabbern: Leyhausen, 1956), a rapid

repetitive biting motion in which the incisors contact the skin or fur, occupies less time than licking.

Nibbling of fur may be accompanied by a simultaneous pulling movement of the head (Durchkammern: Leyhausen, 1956) sometimes apparently in

response to paniculate matter in the hair. The

sides and nape of the neck, the throat, the interscapular region, and the most anterior part of the

neck are not licked and nibbled because of limited

flexibility of the neck. A variety of immobile stances

and reclining postures may be assumed during

either of these activities.

The third type of self-directed behavior that may

be termed bite-pulling involves biting the foreand hindfoot claws with the incisors or the premolars, and pulling backwards with the head. It

appears that either irregularities of the claws or

possibly the claw's hook provides resistance sufficient to create a distinct popping sound as the claw

breaks free from the teeth. The movement undoubtedly removes flakes of keratin from the toenails, but may also dislodge particles from between

the teeth.

HEAD AND NECK.—Rubbing movements of the

head and neck are directed to the forefeet and the
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
NUMBER 239 31

FIGURE 15.—Locomotion and object- and self-oriented contact

patterns in the genet and the Asian palm civet (Paradoxurus):

A, clinging with the everted claws in an infant genet (one

week old); B, head and neck rubbing in a genet; c, penetration of the genet's perineal gland scent-mark about 5 cm into

hardwood; D, scent and urine deposition within a cage

(left arrow: urine encrustation on a vertical cage support;

right arrow: perineal gland scent deposit on the edge of a

diagonal plywood support); E, flank-wiping with a partially

masticated mouse in a female Paradoxurus hermaphroditus;

F, lip-licking in a genet; c, neck scratching in a genet (note

how posture facilitates contact); H, a perineal gland scentmark made by the male Civettictis (arrows point to 1 of 2

streaks made by the glandular lips).

tail in the same manner described for inanimate

objects. Such activity may be interspersed with

other self-directed licking and nibbling movements

and often follows bouts of tail-licking.

NOSE.—Self-directed nasal contact accompanied

by sniffing is most frequently directed to the forefeet, the hindfeet, and the tail, and appears to be

the least frequent of all such behaviors.

HINDLIMBS.—Though the hindlegs can clasp the

tail and can be pressed against the belly if the

animal is suspended by the tail (see "Locomotion

and Its Ontogeny"), most self-directed movement of

the hindlegs involves rhythmic scratching (sichkratzen) characteristic of most mammals. Scratching

is only executed unilaterally in either sitting or

standing positions and orientation to various body

regions is facilitated by a variety of head, neck, and

torso positions (Figure 15G). All body areas can be

scratched with the exception of the forepaws, anterior parts of the forelegs, rump, lower back, belly,

hindlegs, and tail.

FORELIMBS.—The head and the tail are the two

body regions most frequently contacted by the

forelimbs. The paws may be used to draw the tail

closer to the body prior to licking or nibbling, or

may hold the tail in place by standing on it. They

may secure its position as it is licked, nibbled, or

rubbed with the head and neck. An animal lying on

its side or back may also clasp its tail with both

forelimbs. Most frequently, however, the forelimbs

are used in uni- or bilateral wiping of the rostrum,

cheeks, eyes, forehead, ears, and crown. Unilateral

wiping of any part of the head is a relatively simple movement in which a raised forelimb is stroked

forward with an attendant head movement facilitating contact.

Bilateral head-wiping (washing: Eisenberg, 1963;

Gesichtwasche: Diicker, 1965; face and paws:

Ewer, 1967, 1968), on the other hand, involves

coordination of head, tongue, eye, and forelimb movements (Figure 16), and, compared with

other self-directed patterns, usually follows a relatively fixed sequence. The movement consists of

synchronous elliptical motions of the forepaws that

are repeated with stepwise increases in amplitude. Each cycle of forelimb motion begins and is

completed at the mouth where the pronated medial

surfaces of the forepaws come together and are

licked at least once before they execute the next

ascending stroke (Figure 17). During the descending

stroke and possibly during ascent, the head is contacted by the medial surfaces of the manus, wrist,

1 12

FIGURE 16.—Bilateral head-wiping in the genet (animal C;

18 fps): frame 1, licking of a forepaw; frame 12, initiation of

the bout; frames 13-28, wiping of the rostrum and eyes;

frames 224-245, wiping of the ears and crown (note the

pronated forepaws and the role of head and neck movement

during wiping).
32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

e a r

side of head C_

forehea d 11 2 2 2

e y e

 1 1 1 1 1 1 1 1

rostrum • • • 1 I I 1 1 1

ear .

side of head 1

forehead 1 | || |

rostrum ...II I 111

ear ^

side of head l\

forehead _ i OJBVBVB 2

it: ..iiiiiiiiii

4

2 2 2 2 3

2 2 2 3

1 -2^_1 I^ml^m

I1 ? 9 ? J 1 5 7 "=,

I

FIGURE 17.—Sequence of wiping targets and forepaw licking

during bilateral face-wiping in three genets (animals C, I, and

K; each bar represents one cycle of forelimb motion; bar

height corresponds to highest body part wiped; number of

licks directed to forepaws between wipes indicated between

bars; data determined from 8 mm film).

and forearm. Diicker (1957) states that her animals

employed both alternate and synchronous strokes of

the forelimbs. Although in my animals the descending stroke of one limb sometimes traversed a

slightly greater portion of the head, and was hence

more emphatic, only occasionally did one limb lead

the other by as much as perhaps a centimeter, and

never were the limbs seen to be as much as 30° out

of phase during a downstroke.

SELF-ORIENTED CONTACT PATTERNS IN OTHER GENERA

FLANK-WIPING4 (Figure 15E).—This behavior was

frequently seen in the female Pamdoxurus shortly

after she began to masticate dead mice and small

rats. The anterior part of the rodent's body was

transferred from the carnassials to the incisors, and

the wet bolus and hanging hindquarters of the food

was dabbed in several brief posterior strokes on one

or sometimes both flanks; mastication was then resumed. Wiping was only seen as long as the rodent's

body was still at least half intact, and therefore occurred predominantly during the early phases of

feeding on each carcass. The behavior was usually

no longer exhibited after about three mice had been

swallowed. By this time there was often a small

moist patch of fur on each side of the rib cage.

Like neck-sliding, flank-wiping results in the

deposition of a foreign scent on a specific region

of the pelage. In addition, however, a considerable

* I am indebted to Gene Maliniak for bringing this behavior to my attention.

quantity of the animal's own saliva is spread on the

fur. Whether substances other than food are flank

wiped is unknown; hair tonic, though effective in

evoking head and neck rubbing in Genetta, did not

prompt flank-wiping when placed on a paper towel.

ONTOGENY OF SHAKING, STRETCHING,

AND SELF-DIRECTED BEHAVIORS IN Genetta

PRELIMINARY REMARKS.—Shortly after birth headshaking and gaping movements that resembled

yawning could be evoked by tactile stimulation of

the head and neck. At this age stretching movements

consist of simultaneous extension of both limb

pairs and uni- or bilateral extension of the foreand hindlimbs. The movements are often accompanied by neck extension and a concave back

flexion. Stretching during early life is variable and

even after animals are able to walk it usually occurs

in reclining rather than in quadrupedal body positions. The walk-stretch, first seen at 37 days of age,

can probably be performed even earlier.

By two or three days the jaws can execute sustained bites with considerable pressure, and repeated biting movements resembling nibbling are

directed at fingers, blankets, and other soft materials. Incipient nibbling of the body, first seen on

days four and eight (O and K respectively), was

directed to the forefeet; the amplitude of these jaw

movements was noticeably greater than in definitive

adult nibbling. Licking of the nose and lips was also

first seen from four to eight days, and as early as

the eighth day self-directed licking was seen (O and

K).

Animals were first seen scratching themselves in

a sitting position at the ages of 9 and 13 days (O

and Mo). Dttcker (1957) observed that though the

hindfoot was able to execute scratching movements

at the age of 10 days, it was still slightly too short to

reach the body targets until day 15 when the flank,

neck, chin, ear, and mouth were successfully

scratched. The following protocol from my notes

suggests that the apparent inability to scratch with

contact is not necessarily a result of the body's

mechanical limitations.

26 July 1967: Yesterday I was unable to induce baby genets

(2 days old) to scratch themselves. Tonight, continued tickling

of Clawdina's head and earbase with my finger resulted in

the following responses: yawning (3 times), tucking of the

head under the chest (twice), more yawning (3 times). She re-
NUMBER 239

acted to the sixth stimulation by extending the hindleg . . .

forward toward her head, and there the foot quivered very

slightly—seemingly involuntarily.

After a dozen sessions of tickling Ivy's ear, she finally lifted

her hindleg to the side of her head and in a slow . . . movement of that foot brushed her ear lightly. The leg remained

in a partially extended position with the foot lying on the

neck before it recoiled to a flexed resting position.

All the animals I reared could be induced with

repetitive stimulation to scratch either their head or

neck by the age of four days. During repeated stimulation of the ear, the leg often initially scratched

more distal body regions, such as the side and shoulder, and then approached and finally contacted the

stimulated area. The ontogeny of the scratching

response is discussed in more detail in the next section.

Bouts of bilateral face-wiping were first seen performed in extended body reclining positions at 29,

34, 36, and 38 days of age (in I, K, E, and O respectively). The capacity to wipe the side of the face

with one forepaw, however, is usually manifested by

the fourth day provided there is a sufficient outside

stimulus. Diicker's (1957) animals were first seen employing the forefoot for self-contact at the age of 22

days.

Hand-reared animals often may have a scruffy

appearance because they do not respond as the

mother does to presumably unusual or foreign scent

traces on their pelage. Subjectively, it appears that

tactile stimuli induce infants to lick and scratch

themselves.

The finding that localized tactile stimulation is

a simple and effective means of evoking self-directed

mouth and limb movements led to the following

attempt to quantify behavioral responses during

early ontogeny.

MATERIALS AND METHODS.—Animals I and C were

tested a total of 39 and 35 days during a 50-day

period beginning at the age of five days. All tests

were carried out 10 to 60 minutes following

the evening feeding as the animals slept in their

nest box. An effort was made to be consistent in

the method of "tickling" the ear during the course

of testing; however, the possibility of human error

obviously cannot be dismissed. Bouts lasted no

longer than 10 seconds and were usually terminated

sooner when the animal began to respond. An interval of two minutes separated each bout, and each

test comprised from 32 to 41 successive bouts, with

the following exceptions. Animal C was stimulated

11, 20, 26, 24, and 6 times respectively on age days

5, 6, 8, 9, and 51; and animal I was stimulated 5, 6,

21, and 26 times on days 5, 6, 8, and 9. If an animal

did not respond, it was again stimulated two minutes later and the bouts continued until 35 were

completed. Animals at times became active in the

midst of being tested. Tests were continued if they

became quiescent within five minutes; if an animal

remained active longer, testing was resumed a

half hour later. As young animals usually sleep

after being fed, it was hoped that the stimulative

effect of hunger could be controlled by testing

shortly after feeding. Toward the end of the experiment, however, the study animals frequently played

together for up to 40 minutes before going to

sleep. Thus, differing hunger states may have

affected the level of irritability between the beginning and end of the experimental period.

As several different responses could follow a

stimulation bout, responses for each bout were

chronologically numbered and entered onto a

mimeographed sheet divided into 18 labeled response rows and 35 stimulation bout columns.

RESULTS AND CONCLUSIONS.—Ear-twitching and

head-shaking constituted 66.4% of the combined

3717 responses of both animals. In addition, head

jerking movements accounted for 8.0%, and other

motions of the head accounted for 15.8%. Thus,

of all responses, movements of the head and ear predominated (90.2%). Head-shaking and ear-twitching

did not occur with equal frequency throughout

the course of testing (Figure 18G,H). Head-shaking,

often varying over a wide frequency range on

different days, became a less numerous response

by the end of the fourth week. The decline was not

as pronounced for animal C because the initial values were not as high. She was also less viable than

I because of an apparent inability to digest synthetic milk (Esbilac) and did not receive adequate

nourishment between day 15 and day 20 when her

diet was successfully changed. Ear twitching, a less

variable response, showed a distinct linear increase

in frequency of occurrence. The changes in these

two behaviors are not reflections of an altering

overall level of responsiveness, for Figure 18A

shows that the level of the mean number of responses per bout, though oscillating, remained

constant throughout the 50-day period. Frequency

" of occurrence for all behaviors varied widely on

different days, probably as a function of a number
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

of uncontrolled internal and external variables.

Response diversity also varied between days (Figure

18B). The high values of the first four days are

probably exaggerated because of the small numbers

of stimulation bouts, but the elevated figures of the

o 2.0

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5 '5 25 35 45 55

AGE (DAYS)

succeeding four days suggest that responses to

localized tactile stimulation may well be more

varied during early infancy than at later stages.

None of the remaining evoked behaviors contributed more than 1% to the total number of

responses. They can be classified into those that

were distributed (1) sporadically during the entire

50-day period (change body position, twitch legs,

manus to side of head and ear, synchronous forelimb face-wiping, stretching); (2) predominantly

during the first 25 days of testing (extend hindleg,

pes to neck, yawning); and (3) predominantly

during the last 25 days of testing (purring).

Though scratching can be induced by repetitive

stimulation as early as four days after birth, under

these testing conditions it was often manifested in

an inaccurate or incomplete form. An anterior extension of the hindlimb, which normally precedes

scratching, occurred most frequently during the

first 7 to 21 days and only occasionally thereafter

(Figure 18F). Scratching of the neck (Figure 18E)

was irregularly performed by I during the first 20

test days but was an uncommon event in C. While

the ear was understandably the most common target

of scratching (Figure 18D), the side of the head was

occasionally scratched throughout the course of

testing (Figure 18c).

Ear-twitching and head-shaking commonly occurred as part of larger sequences (Figure 19). Table

6 shows that ear-twitching was more frequent overall than head-shaking, more often a single act, and

more common as a serial precedent. In Figure 19 the

following points are apparent. First, of those series

that began with ear-twitching (A) over half (58.8%)

of animal C's and over a quarter (28.5%) of I's responses contained head-shaking (B) as the second

serial element. Secondly, seven dyads alone accounted for the majority (87.2% in I; 80.5% in C)

of all serial combinations that began with earFIGURF 18.—Responses to localized tactile stimulation in two

infant genets (I and C) over a 50-day period (ages 5-55 days;

hollow circles = I, solid circles = C): A, mean number of

responses per stimulation bout (total number of responses

divided by the number of stimulation bouts completed on

that day); B, response diversity (number of response types

divided by the total number of responses per test; because

the first two values for I are .82, they are omitted to conserve space); C-H, percentage of the total number of responses

per test for the following: c, scratch side of head; D, scratch

ear; E, scratch neck; F, extend hindleg; c, shake head; and H,

twitch ear.
NUMBER 239 35

TABLE 6.—Numbers and frequency (percentage in parentheses) of two responses following

bouts of localized tactile stimulation in genets I and C between 5 and 55 days of age

Circumstance

Single act

Preceding act...

Succeeding act..

Total

Ear

393

434

91

917

Animal

twitch

(42.7)

(47.3)

(9.9)

I

Head

165

56

283

504

shake

(32.7)

(11.1)

(56.2)

Ear

299

529

36

864

Animal

twitch

(34.6)

(61.2)

(4.2)

C

Head

76

14

179

269

shake

(28.3)

(5.2)

(66.5)

o

FIGURE 19.—Percentage occurrence in genets I and C of all

behavioral dyads, triads, and tetrads that began with eartwitching as responses to localized tactile stimulation of the

ear (values for I lie below the lines and to the right of the

line between A and B; the total of all the figures for each

animal equals 100%; values between A and the inner ring

of letters represent the percentage occurrences of dyads,

between the inner and the second ring of letters, triads, and

beyond the second ring, tetrads; A = ear twitching, B =

head shaking, c = head jerking, D = move head and replace

in original position, E = change head position, H = wipe

ear with manus, K = scratch ear with pes, N = extend hindleg, o = all other behaviors).

twitching (A). Thirdly, though moving the head (D)

was the most common succedent to ear-twitching in

animal I, on only a few occasions did this pair of

patterns give rise to triads. In both animals, on the

other hand, the sequence of ear-twitching and headshaking (AB) predominantly gave rise to triads terminating with a change of head position (E) and

ear-scratching (K). Data not included in Figure 19

shows that when head-shaking was a dyadic antecedent for animal I (N = 52), the most common

second elements were also change of head position,

26.9%, and ear-scratching, 21.1%. Extending the

hindleg accounted for an additional 13.4%, and of

the remaining 7 dyad types none was more than

5.7% of the total. Animal C displayed only 12 dyads

beginning with a head-shake.

The following conclusions may be made. First,

the most common responses were movements of

the body region most proximal to that of the stimulated area (the ear); these proximal responses subjectively appeared easier to execute than less common self-directed patterns such as scratching. Second, the two predominating movements displayed

somewhat reciprocal frequency changes over the

50-day test period, but they often occurred together

in sequences. The most common triad contained a

low effort antecedent (ear twitch) and moderate

effort succedents (head-shake, ear-scratch, or change

of head position). Third, scratching was the most

complicated type of response and though it was

presumably an effective means of alleviating the

stimulation, it was far less common than proximal

responses (ear-twitch and headshake). Lastly, the

form of the scratching response was sometimes

incomplete and inaccurate. It is difficult to say

whether inaccuracy is a developmental characteristic, because responses to known sources of tactile

stimulation were not observed in adult animals.

The intention movement of scratching (anterior

extension of the hindleg), however, has not in my

recollection been seen during grooming sessions in
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

adults. The pattern is probably an ontogenetic

peculiarity as it was seen in Diicker's (1957) young

animals and in all of those I hand reared. It more

likely results, however, from conditions of the afferent or efferent nervous pathways rather than from

mechanical restrictions of body form.

ELIMINATION

URINATION.—The three squatting postures assumed during perineal marking are also employed

by both sexes during urination, but the crotch does

not contact the substrate. Both hindlegs are commonly on the same level, but if the stream of urine

is directed to low-lying objects one leg may be

cocked over the object or rested upon it. Lateral

swaying of the hindquarters sometimes accompanies

urination in both sexes. As urine is expelled, the

force and volume seems to vary rhythmically, suggesting that flow is controlled by a muscular contraction rhythm in the urethra. In estrous females,

on the other hand, small amounts of urine are often

dribbled upon objects. In both sexes urination is

usually locus-specific, and animals can easily be

trained when young to urinate in a box of litter.

Some genets acquire the habit of urinating in their

water bowl. Once mature, however, they tend to

direct the urine to the elevated edges of the litter

box and other low objects on the cage floor. The

stance is usually stationary, but occasionally, when

urinating on a low-lying object, several short steps

may be taken forward. Animal F was seen to urinate

in this manner on elevated branches, but in the

other animals it was almost always a terrestrial

activity. If frightened or traumatized, however, an

animal will void urine wherever it may be, and, of

course, this often occurs in the inferior animal

during fighting.

DEFECATION.—In adult animals of both sexes

defecation occurs in a squat position with both

legs on the same level. The tail is usually slightly

elevated and may be pumped up and down once

or twice upon completion. The pelvis is also sometimes jerked forward at the termination of the

squat and this presumably coincident with closing

the anal sphincter. In infants, both urination and

defecation can occur spontaneously or can be evoked

in reclining body postures.

ELIMINATION IN OTHER GENERA.—A squatting

quadrupedal stance is assumed (with minor variation) during elimination by Civettictis, Nandinia,

Paguma, Paradoxurus, Hemigalus, and Fossa. Nandinia was frequently seen urinating while slowly

walking forward in a squatted posture. Urination

in Nandinia, and to a lesser extent in Paradoxurus

and Paguma, occurred in and on top of nest boxes,

from elevated branches, and directly on the floor.

Nandinia also defecated indiscriminately about the

cage, but Paradoxurus and Paguma often left feces

on the tops of boxes and shelves. Bartels (1964:198)

reports that in Western Java Paradoxurus excrement is found in elevated positions on top of rocks,

on fallen logs, and "strung out on slim branches . . .

high above ground .. . as well as in collections on

open ground areas." Because "the toddycat keeps

going while relieving itself . . . ," the scats have a

characteristic "strung-out" (rather than heaped)

appearance (1964:198). The same often holds true

for Nandinia. We were unable to train the handreared Fossa to use a litter box, and even when the

animal had free run of a large room it did not display what could be considered locus-specific elimination. The two Civettictis urinated and defecated

in the end of the cage opposite the nest box; the

male frequently urinated and defecated in his water

bowl. In South Cameroun and Senegal, Civettictis

often defecates along trails in small openings devoid

of grass (Gaillard, 1969). Both Hemigalus and

Galidia at the National Zoo urinate in the water

trough.

Urination in the male Paguma and male Civettictis was sometimes closely associated with hindleg

movements. The Paguma, for example, would dribble a small amount of urine on the floor or its nestbox and then scuff its metatarsus in it. On three

occasions during social interaction between the

male and female Civettictis the male was seen

urinating while walking and scratch-kicking. The

urine was spurted to the rear at about 30°-40° to

the horizontal as each hindfoot was alternately

scratched backward with a kicking motion. The

movement resembles the ground-scratching of dogs

except that in the civet it occurs during urination

as the animal walks, and the kicking stroke is

shorter and briefer.

COMMUNICATIVE FUNCTIONS OF ELIMINATION

IN Genetta

Table 5 shows that compared with the various
NUMBER 239 37

types of scent deposits resulting from object-oriented

contact patterns, urine and feces have a strong and

enduring scent value. It is obvious that as odorous

chemical mixtures both substances have an intrinsic

value as media for information about individual

identity, and sexual status and condition (Hediger,

1944). Defecation in the genet is locus specific and

probably communal. Presumably it allows solitary

animals in the wild to detect the existence (through

a chemical symbol) of another animal without confrontation (Mykytowycz, 1969). In captivity both

urine and feces are deposited in a single litter box,

but when two boxes are provided, feces are usually

restricted to one box, while urination often takes

place in both. Anal dragging, a response to voiding

unusually soft feces, probably has only incidental

signal function as it is infrequent and the remaining olfactory trace probably fades rapidly.

Genet urine is apparently chemically distinct

though it has a pungence which characterizes many

viverrine and paradoxurine carnivores. Crawhall

and Segal (1965) found that Genetta tigrina5

excretes unusually large amounts of sulfocysteine,

and concluded that the substance is neither absorbed from the diet nor results from intestinal

microbial metabolism. As amino acids lack the

aromatic properties of cyclic hydrocarbons, it is

doubtful that the compound contributes significantly to the olfactory quality of urine. This

notable feature does suggest, however, that whatever the odorous components in the urine, they

may likewise be distinct and species typical.

SOME INTERGENERIC COMPARISONS

AND DERIVATIONAL SCHEMES

There can be little doubt that body form is a

dominant variable limiting behavioral expression.

Intertaxa behavioral comparisons usually reveal a

small number of fundamental patterns whose

ubiquity probably results from a similar (possibly

primitive) nervous mechanism and a capacity for

movement that is relatively uninfluenced by the

structural divergence exhibited by terrestrial mammalian orders. Definitive adult patterns that fall

into this category are sniffing, licking, and nibbling

movements, scratching with the hindfoot, head and

•These specimens were incorrectly identified by the National Zoological Park (cited Washington Zoo) as Genetta

genetta neumanni.

body shaking, generalized pilo-erection, squatting

eliminative stance, tripedal stance, and the walking

gait. Licking movements (nose, lips, and body),

for example, are generalized motions performed

by the majority of terrestrial species that have

spatulate intra-oral tongues (Type I tongues of

Dor an and Baggett, 1971).

Though coordination of fundamental patterns is

usually similar between species, differences are often

clear reflections of structural adaptations for speed

or arboreality. Both Fossa and Nandinia, for instance, can touch the pes to the inside of the

opposite leg, but this scratching orientation is less

commonly displayed by the former than the latter.

The lack of an extensive rotatory capacity in Fossa's

hindlegs may be among the factors accounting for

this difference. The dexterity of Nandinia's hind

leg on the other hand probably reflects a primary

adaptation for arboreal locomotion, especially headfirst vertical descent. As adaptations for climbing

and speed imbue differing mechanical properties

to the skeleton and muscles, it is not surprising that

behavioral differences between species do not always

parallel taxonomic distinctions. This contention is

supported by Table 7, comparing the distribution

of several behavior patterns between members of

four subfamilies. I have included the first two

behavioral items in Table 7 (bipedal stance and

headfirst vertical descent) to give an indirect comparative measure of arboreal specialization. The

assumption is that civets, unable to descend trunks

headfirst or stand bipedally, exhibit morphological

correlates of at least incipient cursorial specialization, namely digitigrady and suppression of transverse and rotatory limb motion (Howell, 1965). In

the following discussion, based in part on Table 7,

similarities and differences should be viewed both

in terms of locomotory adaptation and systematic

status.

Both the long- and walk-stretch are probably the

most common and frequently performed stretching

patterns in carnivores as well as many other mammals. In their complete form, both of these patterns

consist of two primary and independently performed components, namely stretching of the forequarters and stretching of the hindquarters. The

differences between the patterns result from the

association of the walk-stretch with forward progression. In the typical walk-stretch the forelegs

are not uniformly extended as the forequarters are
38

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 7.—Comparisons of behavior among selected viverrid genera ( + = present, — — absent,

+ ? or — ? = probably present or absent but substantiation needed, blank = uncertain, a =

personal observation, b = Vosseler 1929, c = Ducker 1957, d = Huf 1965, e = Diicker 1971,

f = Albignac 1970a, g = Albignac 1970b)

Stationary stances

Bipedal stance..

Head first

vertical

descent

Stretching

Long-stretch

Walk-stretch

Arched-back

stretch

Object-oriented

contact patterns

Metatarsus

scuffing

Head and neck

rubbing

Neck sliding

Scent-marking

patterns

Quadrupedal

squat

Perineal drag...

Normal

quadrupedal...

Reversed upright

quadrupedal...

Upright

quadrupedal...

Self-oriented

movements

Bilateral head

wiping

References

Viverrinae Paradoxurinae Hemigalinae

Cryptoproctinae

a a ac ad a ae

_?

afg

depressed and the hindlegs take one to several stifflegged steps. In the typical long-stretch, the forelegs

are equally extended in the first phase, and the

hindlegs equally extended in the second phase.

The variation of these patterns within a species

or individual, however, does not suggest a clear

evolutionarily interpretable trend. Genetta, for example, performs walk-stretches with the forelegs

equally or subequally extended, and partial walkstretches in which variation in forelimb positions
NUMBER 239 39

are manifested without the hindquarter stretch

(stiff-legged walk). The same holds true for Nandinia and Paradoxurns. Civettictis, on the other

hand, stretches like a dog: the forebody is depressed

as the forelegs are equally extended forward. Nevertheless, both the civet and the domestic dog occasionally terminate such a forequarter stretch with

several short steps of the posteriorly extended

hindlegs. Quantitative data are clearly needed to

establish relative frequencies of the different

stretching coordinations.

The categorization of complicated and component coordinations (such as fore- and hindquarter

stretches) into walk- and long-stretches obscures the

possibility that these component coordinations were

the precursor elements that combined in time to

produce the walk- and long-stretch. Alternatively

it is possible that the widespread walk-stretch pattern was the prototype that, through fragmentation and changes in form, gave rise to fore- and

hindquarter stretches, and evolved into the longstretch through loss of the association with walking.

The arched-back stretch is equally enigmatic, occurring in erinaceids (Poduschka, 1969) as well as felids

and viverrids.

Bilateral face-wiping is an almost identical movement in the Viverrinae and Paradoxurinae exhibiting it (Table 7); Nandinia, which differs slightly

from Genetta, holds its forepaws in a semiprotracted

position and consequently after each descending

stroke they do not meet on their medial surfaces.

According to Diicker (1965), the pattern in Viverricula also resembles that of Genetta, except that

it is performed in a reclining position. Civettictis

on the other hand lacks this behavior, though I

once saw the female make strained but synchronous

forepaw contact to the nose as she reclined in a

lateral extended position. The ability to touch the

head simultaneously with both forepaws, however,

is widespread within the Carnivora; even domestic

dogs will eventually remove a rubber band placed

about the snout with simultaneous use of the dew

claws. Though bilateral face wiping has not been

seen in paradoxurines other than Nandinia, the

forelimbs appear to have ample dexterity. Unilateral face wiping, on the other hand, was uncommonly seen in Paradoxurus, Paguma, and Arctictis,

and rarely seen in Civettictis and Fossa.

With the exception of Burger's study (1959),

bilateral face-wiping has been overlooked as one

of the most complex and stereotyped of mammalian

behaviors. The resemblance in the coordination

and form of this pattern between Insectivores,

Tupaiids, rodents, marsupials, and carnivores

implies either remarkable convergence in neurophysiological evolution, or retention among disjunct

lineages of a primitive and formerly widespread

pattern. The genera I have observed6 exhibited

these similarities: (1) bipedal stance, (2) forelimb

motions with progressively increasing amplitude,

and (3) forepaw licking strongly linked to the completion of each wiping motion (at least initially during the bout). The primary differences between

species relate to the posture and action of the

manus, the phase relationships of the forelimbs to

one another during wiping, and the speed with

which the movement is executed. No combination

of behavioral, morphological, or ecological criteria

can be used to predict accurately which species

possess bilateral face wiping within their repertoires,

and consequently it has little utility as a taxonomic

character.

Judging subjectively, the pattern seems to occur

only in species with either (1) lax pelage, or (2) a

tactile receptive field on the face composed of long

vibrissae and possibly scattered tylotrichs, and (3)

a body form sufficiently generalized that the limbs

have considerable freedom of movement in several

planes. This grooming ability is displayed by nonbrachiating arboreal and semiarboreal species, and

those terrestrial species having plantigrade feet. The

existence of the pattern in carnivores is highly

unusual and suggestive of a more widespread occurrence among the extinct viverrines that presumably

gave rise to modern forms. This is confirmed by the

occurrence of unilateral face-wiping in many felids

(Leyhausen, 1956) and its resemblance to bilateral

wiping in the form of the motion and the alternating lick and wipe pattern. Furthermore, no

movements resembling bilateral face-wiping are

known in the most convergently similar arctoid

carnivores, such as Potos and Bassaricyon (PoglayenNeuwall, 1962, 1965) and Bassariscus (H.

•Viverridae: Nandinia, Genetta, Prionodon; Didelphidae:

Didelphis virginianus, Marmosa robinsoni, Caluromysiops irrupta, Caluromys lanatus, Chironectes minimus; Phalangeridae: Phalanger gymnotus, P. orientalis, Petaunis breviceps;

Dasyuridae: Dasyuroides byrnei, Sminthopsis macrura; Insectivora: Sorex sp., Blarina brevicauda, Microgale talazaci,

M. dobsoni, Echinosorex gymnurus.
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Egoscue, pers. comm.). On the other hand, this

behavior is exhibited by several mustelids (Mustela

erminea, M. frenata, Spilogale putoriw: Egoscue,

pers. comm.). The bilateral forepaw patting motion

that Enhydra lutra directs to its head superficially

resembles face wiping, but because the movement

lacks an association with paw licking and is directed

to the entire body, it probably derives from a behavior pattern other than the one in question.

In the felids and viverrids where uni- or bilateral

face-wiping is a regular constituent of the toilet

that precedes activity or follows feeding, the pattern's appearance seems to be largely self-generated,

or at least responsive to afferent input from the

stomach or the mouth, lips, and vibrissae. Ewer

(1967) has postulated a scheme which simulates the

sequential and repetitive character of grooming

sessions based on the interaction of the excitatory

levels of the component actions. In those species

which only on occasion employ unilateral face

wiping, its appearance seems largely governed by

external stimuli, and the causal linkage of unilateral wiping to other self-directed patterns seems

to be weak or lacking. Fossa fossa, for example, was

rarely seen to wipe its snout, but a loose, deciduous

premolar precipitated a prolonged series of wiping

motions that eventually dislodged the tooth.

Head- and neck-rubbing, and neck- and shouldersliding (Table 7) are basically similar in that largely

overlapping anterior parts of the body are rubbed.

The former pattern, however, is generalized in the

sense that it is evoked by a variety of circumstances

and its component motions exhibit variable forms,

duration, and temporal organization. On the contrary, neck and shoulder sliding seems to be evoked

only by certain aromatic substances and consists

of one motion directed to alternate sides of the

head and neck. Though it is tempting to interpret

neck and shoulder sliding as a simplified and

exaggerated version of Genetta'% rubbing movement,

two observations suggest a more logical alternative:

(1) when thoroughly wet, domestic dogs, mustelids

(M. frenata, M. erminea: pers. observation), and

viverrids (Genetta, Nandinia, Fossa) sometimes propel themselves forward with the hindlegs while

sliding on the side of the head, neck, and, to a

lesser extent, the shoulder; (2) sliding movements,

in which substrate contact coincides largely with

the extension rather than retraction phase of the

neck motion, occur in several species either as a

response to a strong odor field (canids, Eira barbara,

the three viverrids listed above), and/or in species

that have a dermal scent-producing gland in the

cheek, throat, neck, or sternal region {Trichosurus

vulpecula, Helogale undulata: Zannier 1965, probably Herpestes auropunctatus: D. Nellis, pers.

comm.). In dogs neck-sliding directed to animal

dung may be repeated several times on alternate

sides of the neck, or it may occur only as a brief

preliminary to rolling about on the sides and back.

When in heat, female domestic cats slowly glide the

neck to the shoulder on the substrate while standing

(Leyhausen, 1960). This behavior is apparently

spontaneous.

In comparison, neck and shoulder sliding as a

prefatory component of body rolling and rubbing

appears to be an incipient expression of the definitive pattern seen in civets. According to Diicker

(1957) neck rubbing (Kinn- und Halsscheuern) in

Viverricula is followed by vigorous bouts of body

rolling and rubbing. The factors that have selected

for the abandonment of body rolling in favor of

only neck sliding are obscure; however, the simplification of the movement may be linked to intensification of the sniffing role; brief sniffing bouts are frequently interposed between each sliding motion

during the onset of this activity.

Considering other patterns of object-oriented

contact, Table 7 attests to the intra- and interspecific variation in the methods by which the

selected viverrids appress the pregenital or perineal

gland to inanimate objects. The most common

marking methods, namely the perineal drag and

quadrupedal squat, have a distinct resemblance to

anal dragging and differ from it chiefly in the angle

of the pelvis and so the emphasis of contact. The

physical similarity of these three movements, and

the fact that anal dragging is a prevalent response

among morphologically generalized mammals to

moisture, feces, or paniculate matter on the anus

or hindquarters, suggests that both perineal and

anal dragging primitively shared common causal

features related to elimination. (For the sake of

clarity in this discussion, the terms anal drag and

perineal drag will be used to designate respectively

the probable primitive cleaning movement and the

scent deposition movement assumed to be its derivative.)

The hypothesized significance of autonomic

elimination for the evolution of many types of
NUMBER 239 41

anogenital scent glands is founded on the following

observations. In many mammals reflex discharge

of urine and feces is most frequently caused by

hostile inter- and intraspecific interaction, often

resulting in anogenital contamination and consequently followed by anal dragging and self-directed

licking and nibbling. Contained within the autonomic elimination reflex and the circumstances of

its occurrence are behavioral "building blocks" or

preadaptations for specialized forms of scentmarking. The building blocks include two distinct

modes of scent deposition (urine and feces), and a

potential social referent and/or message receiver

(the animal evoking the autonomic response). To

elucidate, both autonomic defecation and evacuation of the anal gland share common features of

agonistic contexts and protrusion and eversion of

the anus. The implication is that anal protrusion

was a possible source of anal scent emission. Anal

dragging may likewise have served as a behavioral

vehicle that potentiated the development of glandular fields in the anal, perineal, and pregenital

regions.

The remaining three viverrid marking patterns

(Table 7; Figure 20) differ from the perineal drag

in that the association with forward progression has

been relinquished and glandular deposition is mediated by a simple pressing motion (quadrupedal

stance in Civettictis), a uni-directional wipe (reversed upright quadrupedal stance in Genetta), or

a rubbing motion (antero-posterior in Cryptoprocta,

or lateral in Genetta). A fundamental similarity of

the two upright quadrupedal patterns (reversed

and normal) to the perineal drag can be appreciated

by visualizing the necessary adjustments in posture

that would accompany rotation of the marking

log seen in Figure 20 to a horizontal plane. The

implication that both patterns arose as concomitant

expressions of arboreal locomotory abilities need

not be true. The postural properties of the reversed quadrupedal upright position, for example,

characterize two situations that could conceivably

have served as transitional activities as denned by

Lind (1959). As an animal steps from an elevated

position to a lower level, the effective angle of

descent and center of gravity are decreased by

depression of the hindquarters and the acquisition

of a spinal concavity (Figure 20). The loss of this hypothetical stepping-down component, which would

convert the pattern to its definitive form, could

POSSIBLE PRIMITIVE SOURCES:

NORMAL AND AUTONOMIC

ELIMINATION & ASSOCIATED

ANAL-PERINEAL DRAGGING

gland origins

(anal, perineal. & preputial)

GLAND DRAGGING

TRI-&

QUADRUPEDAL

SQUAT

(loss of

locomotory

component)

FIGURE 20.—A derivational scheme of some stances employed

by viverrid carnivores during scent-marking (arrows point to

specific stances exhibited by extant species and a hypothetical

intermediate stage; black arrow = logical evolutionary pathways of stances in which glandular contact with the substrate

is mediated by variations on the squatting theme; white

arrow = two possible (but rather tentative) routes for the

development of the normal quadrupedal marking stance from

a squatting stance).
42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

have occurred through an increased dependence

upon olfactory releasers and an escape from the

transitional action. The transitional act hypothesis

is supported by the observation that in Cryptoprocta the upright marking stance is frequently

preceded by substrate clawing accompanied by a

lordotic spinal curvature; this pattern may have

likewise undergone similar changes.

Alternatively and perhaps more feasibly, marking

in the reversed upright quadrupedal stance may

have evolved as an exaggeration of the tendency to

place a slightly elevated mark by raising a hindleg

over or by resting it upon the marking object

(e.g., Genetta tigrina, Suricata suricatta, Herpestes

edwardsi: Ewer, 1968). Cricetomys gambianus (Ewer,

1967) and female Speothos venaticus (Kleiman,

1966) achieve more elevated marks by walking backwards up inclined surfaces; such a method is feasibly intermediate to the technique used by Genetta

in which the hindquarters are raised up to the

marking position via a quick handstand rather than

climbing up backwards.

The simple pressing glandular contact and the

absence of the squat component indicate a minimal

resemblance of the quadrupedal marking pattern

to the perineal drag. Derivation from the perineal

drag or squat presumably involved a concurrent

posterior shift in glandular position for, other than

a slight spreading of the hindlegs and lifting of the

tail, the quadrupedal marking pattern exhibits no

other postural adjustments. De novo origin of this

pattern is tenable only if it is assumed that perineal

glands evolved independently in the semiarboreal

and terrestrial viverrines after divergence from the

common ancestor. The remote resemblance of

marking with the perineal gland in a normal

quadrupedal stance is perhaps a manifestation of

Civettictis' particular subcursorial habitus.

Behavior of Associated Animals

Patterns of behavior performed by solitary animals also occur in the presence of other animals.

A number of behaviors, however, seem to occur

predominantly in the presence of conspecifics because their execution requires an animal referent,

or because stimuli and contexts associated with

interaction are usually absent in solitary situations.

The wisdom of classifying some of these behavior

patterns as either solitary or social is debatable.

Generalized pilo-erection, for example, is mediated

through any or all of the sensory systems; the

characteristics of the stimuli are not specifically

limited to social contexts. Under the conditions of

this study, though, certain patterns of pilo-erection

were seen primarily in social contexts, hence their

inclusion in this section.

The descriptions in the first part of this study

defined types of body motion as they were identified

and recorded during social encounters and subsequently analyzed. With the exception of vocalization that is distinguished by broadcast transmission,

the behaviors in this section are classified on the

basis of orientation and the spatial characteristics

of their occurrence. It is assumed that these patterns

have a communication function. Both the perceptual effect and, presumably, the message of each

pattern is or may be modified when executed concurrently with other patterns; thus, the behaviors

described under each category represent components

that may be combined within and between categories and modified by independent variables such

as velocity of movement and posture. The second

part of this section analyzes the organization of

signals in time and the dynamics of social interaction in Genetta and Civettictis.

COMPANION-ORIENTED LOCOMOTION

An animal's locomotion is often affected by the

presence of another animal. The criteria used to

distinguish different types of companion-oriented

locomotion are listed below.

APPROACHING.—Technically, any movement of

one animal that decreases the distance between it

and a companion may be defined as an approach.

Such a definition would include movement tangential to, but not specifically oriented to the other

animal. As any movement of one or two animals

will involve spatial changes that may have no immediate social significance, the term approach is

used herein only to describe those cases in which a

decreasing distance between two animals is accompanied by at least one animal maintaining a frontal

orientation to some part of the other animal's

body (Figure 23A).

DEPARTURE.—The term is used to describe the

net movement of one or both animals away from

the immediate vicinity of the companion. In most

cases a departure is defined as the separation of
NUMBER 239 43

animals engaging in contact, or the movement of

one animal out of a one body length perimeter of

space surrounding the companion's head and body.

FOLLOWING.—An animal is described as following

when its locomotion and spatial orientation are

closely correlated with and apparently guided by

the companion's movements. Following is usually

a distinct action with the animals trotting briskly

in close proximity, the follower at the rear or side

of the leader.

CIRCLING.—When the locomotion of each of two

animals is oriented toward the side or hindquarters

of the partner, the course of movement is circular.

This is seen during mutual sniffing of the hindquarters or when both animals attempt nasal contact with the partner's rear. This definition of

mutual orientation to the companion's body

excludes following in which the path may also be

circular.

ENCIRCLING.—Occasionally as one animal remains

stationary or moves slowly, its companion walks or

trots about it in a tight circle.

JUMPING OVER.—Though this pattern may involve an approach and departure, it is best described

as a separate case in which the body of one animal

passes over the companion's body. During bouts of

interaction involving extensive body contact, an

animal may jump over another without leaving the

partner's proximity. It may also be executed by an

animal traversing a tree limb, in order to negotiate

the obstacle presented by a resting companion's

body.

GAIT AND POSTURAL VARIABLES.—Speed of movement and postural variants of the walking gait add

additional though somewhat overlapping dimensions to simple spatial changes resulting from the

locomotory movement of two animals (see "Gaitassociated Postures" under "Locomotion and Its

Ontogeny"). While approaching, following, and

departure may involve any gait, circling and encircling are performed only in a walk or trot. The

association of postural configurations with types

of companion-oriented locomotion has not been

consistently pursued, but it appears that nearly all

the variants diagrammed in Figure 5B-F are exhibited by approaching animals, while departing

animals rarely exhibit postural variants involving

crouched forequarters (Figure 5B,D). When a directed gait is interrupted by a stance, the postural

configuration of the spine is usually retained. As

TABLE 8.—Associations of postural variants (companionoriented stances of tripedal and quadrupedal types, characterized by protracted visual fixation) and torso orientation to companion in Genetta tigrina (+ + = most frequent

association, + = less frequent association, — = uncommon

association)

Postural variant

Full Crouch

Normal

Crouched fore- and

semi-raised hindquarters

Semi-raised foreand hindquarters.

Maximum raised

fore- and hindquarters

Frontal Tangential Lateral

there is no discrete distinction between a slow walk

and brief alternating tri- and quadrupedal stances,

the stances described in the following section

represent the stationary extreme of a graded contiuum of noncontact-oriented behavior.

COMPANION-ORIENTED STANCES

Visual fixation of a companion requires only

that the head be oriented in the companion's direction, but in a number of circumstances, particularly

when a stranger is encountered, relatively long

periods of time may be occupied by staring. The

stances assumed during visual fixation' are characterized by (1) rigid-appearing immobility, (2) interruption by brief periods of walking but often

unbroken visual contact, and (3) the maintenance

of a particular body orientation to the companion

irrespective of the companion's movement. Table

8 presents a rough approximation of the way these

stationary body positions are associated with the

orientation of the torso relative to the companion's

body. It is clear that with the exception of the

normal quadrupedal stance there is a tendency for

the body to be low during frontal orientation and

high during lateral or tangential orientation (Figure 23A). This is in part a result of the association

of the low or crouched stances with approach

toward the companion whereas the high or archedback stances are usually seen in a context of

avoidance.
44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

12

FIGURE 21.—Bipedal pivot of the African palm civet (Nandinia) based on tracings of 16 mm film (24 fps), in which the

animal took several bipedal steps.

The bipedal stance and the elongate quadrupedal

crouch are usually maintained for relatively brief

periods, the latter being associated with approaching a companion and establishing nasal

contact.

SIMILAR STANCES IN OTHER GENERA.—Civettictis,

Paradoxurus, Paguma, Nandinia, and Fossa are

capable of assuming a crouched posture while stationary or walking. Though not all these genera

display equal facility in maintaining this position,

crouching is fairly widespread among civets. With

the exception of Paguma, the posture involving

semicrouched forequarters and semiraised hindquarters (Figure 5D) is also seen in these genera as

a companion-oriented stance. This body position

can be converted to a low-intensity arched-back

posture by straightening the forelegs and slightly

arching the back. This has been seen in Paradoxurus, Nandinia, and Fossa.

In the male Nandinia, this half crouched, half

raised body carriage was associated with slow and

deliberate walking on branches, accompanied by

visual fixation of the observer or another animal.

The male would reverse the direction of his movement by rising "stiff bodied" into a bipedal position

and pivoting on his hindlegs. Eye contact with the

other animal during this maneuver usually was

not broken. An unusual form of this bipedal pivot

was displayed by the male toward a strange female

(Figure 21). The animal quickly approached the

female, rose into a high bipedal position, and

pivoted to one side while holding the forelegs up

and tightly flexed to the sides of the body. It can

be seen in Figure 21 that the animal actually took

several steps while standing upright.

Intense arched-back postures have been seen only

in Fossa. I am reasonably certain that it does not

occur in Civettictis and Nandinia, but neither

Paradoxurus nor Paguma was seen under conditions

conducive to this behavior (e.g., fighting). Intense

arching in Fossa occurs as a laterally oriented stationary stance, and the tail may be held vertically

erect.

In Civettictis tangential or lateral orientation of

the head, neck, and body to the companion is an

outstanding aspect of social interaction. There is

no simple way this can be described because the

position of the head and neck is clearly dependent

to a large extent upon the attitude, proximity,

and orientation of the companion. In general, the

neck is held at the horizontal or slanted downwards while the head is maintained in the same

plane (Figure 22A) or deflected downwards to as

much as 60° (Figure 22B). The head is usually

rotated slightly upwards in the partner's direction

(Figure 22A,D) and may be turned slightly toward

or away from and out of the companion's sight.

Though it is possible for an animal to display a

lateral head and neck orientation to a companion

while the trunk has a frontal attitude (Figure 22B),

it is more common for the trunk axis to be tangential or lateral to the companion's head and

neck (Figure 22A,C,D). Tangential or lateral body

or head and neck orientations to a partner occur

during slow walking in a normal posture, in a

normal or crouched quadrupedal stance (Figure

22c), or in reclining positions (Figure 22A,D).

NONCONTACT BODY MOVEMENTS

ERRATIC HEAD MOVEMENTS.—The simplest form

of this movement involves a rapid vertical bobbing

motion of the head with the neck contributing

minimally. This is most frequently seen when an

animal is approached or followed. Such erratic

motions result from a rapid twisting of the neck,

and lateral, circular, and vertical jerking movements of the head, often associated with a brisk

trotting gait, or brief bipedal stances.

ALTERNATING EAR MOTIONS.—As described under
NUMBER 259 45

FIGURE 22.—Lateral and tangential orientations of the head, neck, and body in the African

civet (Civettictis): A, head and neck deflection in a reclining animal; B, lateral head and

neck orientation by an approaching animal (male); c, assumption of a reclining position by an

animal receiving head-darts; D, head deflection by a recumbent animal towards the superior

companion.

the primary senses, this behavior may be directed

toward another animal or to an area of interest.

It is difficult to discern whether the movement itself

or the independent perception of sound triggers the

same response in companion animals.

TURNING TOWARD.—Any motion of the body or

its parts resulting in a frontal orientation of the

face toward the companion is denned as turning

towards. Facial exposure can be achieved by either

turning the neck or pivoting the entire body in the

companion's direction. This pattern may initiate a

long period of visual fixation accompanied by a

companion-oriented stance or it may involve only

a brief gaze in the other animal's direction.

TURNING AWAY.—Any movement that terminates

facial exposure to a companion is denned as turning

away.

TAIL WIGGLING.—An erratic and rapid bending

and extension of the tail along its length produces a

spasmodic jerking motion that in its most intense

form produces a rattling sound against wooden

floors or a rustling sound in leaf-litter substrates.

In my animals, tail wiggling was always performed

in either a ventral retracted limb or an extended
46

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

body reclining position, but according to Leyhausen

(pers. comm.) a violent thrashing motion of the tail

accompanies arched-back rubbing in Genetta

riippelli.

ANAL GLAND SCENT EMISSION.—The emission of

anal gland scent is associated with fighting, intense

attempts to avoid or escape from another animal or

painful stimuli. Hence, it is not accompanied by

any particular stance or movement. On the one

occasion I observed scent emission at close range,

the anus was briefly opened several millimeters and

the fluid ejected as a barrage of about two dozen

small drops that traveled about 2 decimeters before

striking a surface. The force of the emission is

weak, and the associated circumstances are such

that scent is usually deposited close to the interacting animals. The scent almost always contaminates the fur of the tail and that surrounding the

anus; animals usually spend considerable time

licking these regions following scent release.

Anal gland scent appears to arise from two

spherical structures lying beneath the tail on either

side of the anus. The scent is a thin translucent

coffee-colored fluid that is highly volatile and has

a slightly noxious musky odor. The odor lingers in

the air for several minutes after release; to the

human nose it is detectable within a radius of about

3-4 meters in a room with minimal air movement.

Even after several weeks the scent residue emits a

mild musky but not unpleasant odor that is detectable at close range; washing with detergent and

water will not completely remove it from clothing.

PILO-ERECTION PATTERNS.—The longer hair on

the head, trunk, upper legs, and tail is erected in

a variety of circumstances that are not always related

to the presence of another animal. The degree of

erection is variable and not necessarily uniform on

the body surface. Disturbance of the nap of a

pelage often occurs during scratching and objectoriented contact, and uniform erection of the hair

seen in resting animals is presumably associated

with thermoregulation. In addition, certain patterns

of pilo-erection are exhibited in circumstances involving other animals.

Vibrissae: Movement of all the facial vibrissae

from depressed to an erect position is presumably

under the control of pili-arrectores muscles and

superficial facial musculature. Maximal mobility

is displayed by the mystacial vibrissae. The anterior

movement of these hairs, whether or not associated

with gaping of the jaws, always involves a squaring

of the rostral contour (Figure 26). Erection and

depression can be either gradual (and arrested at a

certain degree of erection) or sudden. Sudden

changes in vibrissae position (<1 sec) seem to be

maximal changes involving erection. Depression is

usually a gradual process. The mobility of the

other facial vibrissae seems to be less developed

and less noticeable.

Spinal Crest Hair: The black hair of the spinal

crest, extending from immediately behind the

shoulder to the base of the tail, slightly exceeds

the length of the surrounding hair (Table 9). Thus,

when there is uniform erection of hair on the torso,

the crest is slightly elevated (Figure 23B); moreover, it appears that the spinal crest itself can be

TABLE 9.—Average hair lengths (in mm; range in parentheses) of spinal crest and tail in a

sample of 18 Genetta tigrina skins in NMNH (sides = 3 cm from dorsal midline; tail base

= third white tail ring to rump; tail tip = distance from end equal to tail base distance

from rump)

99 dorsum

side

diff.

dtf dorsum

side

diff.

99

 + oV dorsum

side

diff.

Shoulder

17.7

17.1

0.6

19.2

18.8

0.4

18.4

17.9

0.5

(16-20)

(16-19)

(0- 2)

(16-22)

(16-23)

(-2- 4)

(16-22)

(16-23)

(-2- 4)

Spina1 Crest

Midback

21.7

19.4

2.3

24.0

18.8

5.2

22.8

19.1

3.7

(20-24)

(19-22)

(0- 4)

(20-31)

(17-24)

(0- 9)

(20-31)

(17-24)

(0- 9)

Rump

25.8

18.8

7.0

26.1

20.1

6.0

25.9

19.4

6.5

(22-30)

(16-20)

(5- 9)

(22-31)

(17-24)

(2-11)

(22-31)

(16-24)

(2-11)

24.8

24.8

0

25.5

25.5

0

25.1

25.1

0

Tail

Base

(21-27)

(21-27)

(22-29)

(22-29)

(21-29)

(21-29)

20.4

20.4

0

21.6

21.6

0

21.0

21.0

0

Tip

(16-23)

(16-23)

(16-26)

(16-26)

(16-26)

(16-26)
NUMBER 239 47

raised to a greater extent than surrounding hair.

Maximal and prolonged erection of the crest,

however, seems to be linked with a uniform level of

erection in the torso hair. When the body hair is

depressed, there is no indication of a crest.

Tail Hair: The degree of erection may be uniform or may vary along the length of the tail. A

uniform level of erection seems to be the most common condition (Figure 23c). Lengthwise piloerection gradients are usually unidirectional with

either the terminal black segment or the first 2-4

white rings being the expanded portions. The

velocity of hair erection and compression (hair flux)

varies from moderately rapid (<2 sec) to gradual,

but cine film is necessary for accurate description of

this aspect. While gradual expansion of the tail

hair is more or less continuous, rapid expansion or

compression is often, perhaps most frequently,

stepwise. That is, rapid flux in erection or compression is followed by a period of very slow change

in either direction.

NONCONTACT MOVEMENTS IN OTHER GENERA.

With the exception of alternating ear motions and

tail wiggling, Civettictis exhibits all the noncontact

body movements described for Genetta. The African civet is reputed to expel an anal scent (Walker,

1968), but though my animals fought on several

occasions this was not detected. Unlike Genetta,

the hair on the tail is not erected in stepwise increments. This pattern of hair flux is seen, however,

in the well-developed spinal crest. Erection of the

crest results in an obvious increase in lateral body

dimension and adds a light-colored stripe to the

contour of the back above the level of the spine;

this is not visible when the crest is relaxed because

the light-colored basal segments of the hair are

covered by the longer black distal segments.

COMPANION-ORIENTED CONTACT PATTERNS

Head

SNIFFING (NASAL CONTACT).—Sniffing of another

animal does not necessarily involve contact, but if

not repelled an animal will almost always bring its

nose into contact with the fur of its companion.

The nose may lightly contact the tips of the companion's hair or may be pushed through the hair to

the skin. The rhinarium, lips, and the lightly furred

zone between them are often rubbed through the

underhair and against the skin (Figure 23D). A

variety of facial expressions may be displayed as an

animal sniffs and contacts a companion with its

nose. When sniffing without contact the eyes are

usually wide open and the ears erect and directed

forward. If nasal contact is involved, there is a

tendency for the eyes to be squinted and the ears

oriented laterally (Figure 23D). When the perineal

gland, vagina, urine, or an estrous female are

smelled and/or licked by a male, the upper lips

may be lifted slightly and the corners of the mouth

retracted to expose the canines and premolars.

This activity, Flehmen, is presumably associated

with olfactory and gustatory perception via Jacobsen's organ. The mouth is usually slightly opened,

the eyes squinted, vibrissae retracted, and the orbits

of the ears directed laterally in a semierect position.

LICKING AND NIBBLING.—As described for the

solitary animal, these patterns may also be directed

to a companion. Both types of mouth contact are

usually associated with a relaxed facial expression.

The eyes may be squinted or closed and the ears

oriented laterally in a semierect position.

BITING.—Duration of contact, force of compression, and head movements during contact are three

biting parameters that probably form a graded

continuum. Mouthing occurs when an animal's jaws

engage with another animal's body but there is

little or no compression (Figure 23E); biting occurs

if the force is strong. Duration of biting and

mouthing is highly variable. Facial characteristics

of hard biting include closed eyes, depressed mystacial vibrissae, and laterally directed ear orbits that

are folded posteriorly above the notch. During soft

biting or mouthing the eyes may be wide open or

squinted; the ears are usually held with the orbits

oriented laterally, but they are not folded above

the notch (Figure 23E).

HEAD-DARTING (Figure 24).—This is a highly variable movement consisting of several components

and involving several levels of integration. It is a

rapid form of head extension which may or may

not be followed immediately after by head retraction. It may be executed with a closed mouth,

with an open mouth, or with an open mouth and

an explosive hiss emitted at the end of the extension phase. Figure 25 displays the variability in the

duration of extension and retraction phases and

the way an open mouth position may be associated

with this activity. Thus, head-darting may or may
48 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 23.—Patterns of companion-oriented locomotion, movement, and contact in the genet:

A, crouching animal approaching a companion exhibiting an arched-back stance; B, low intensity arched-back stance exhibiting general pilo-erection and the spinal crest; c, maximal

pilo-erection of the tail; D, juvenile genet sniffing the spinal crest hair of another animal

(note facial expression); E, juvenile animal mouthing its companion's tail; F, pawing by an

animal engaged in wrestling.
NUMBER 239 49

1.3 4

1 3 4

5 8 10

10 8

FIGURE 24.—Extension and retraction phases of head-darting

in the genet (animals E and Fr; based on 16 mm film, 24 fps;

shaded images represent last frame in series; in this sequence

recipient began to respond in less than i/A second).

EXTENSION/RETRACTION

FRAMES

12 10 8 4 0 4 8 10 12

FIGURE 25.—Duration of extension and retraction phases in

eight samples of genet head-darting recorded on 16 mm film

(24 fps) of E head-darting at Fr (none involved contact with

Fr and the occurrence of hissing is not indicated; black

bars = extension and retraction of the head, clear bars =

opening of the mouth).

not involve contact with a companion. When it

does involve contact, the companion is most commonly struck with the incisors and canines of the

partly opened jaws. Brief snapping bites are delivered less often. Associated with such contact is

a momentary squinting or closing of the eyes (Figure 24). A backward deflection of the ears is a far

less frequently associated change. The mystacial

vibrissae are often extended anteriorly preceding a

head-dart and during the early part of neck extension, but as the subject is approached, they are

retracted at least momentarily. The vibrissae may

retain an extended position if the velocity of the

extension phase is relatively slow.

JAW-GAPING STARE.—The mouth may be gaped at

20°-30° for varying amounts of time as an animal

stares at a companion. Extended vibrissae are

usually associated with a widely gaping jaw (ca. 40°)

and a strong tendency to head-dart at a companion.

The mouth may be opened briefly and repeatedly

while the animal is staring and as it inhales between

growls.

ERRATIC HEAD MOVEMENTS (see "Noncontact

Body Movements").—When executed in the close

proximity of another animal virtually any part of

the head or neck may touch the companion's body.

Limbs

Three kinds of forelimb movement directed toward other animals can be discerned. Whether involving contact or not, pawing motions are usually

brief and similar in form to those directed at inanimate objects (Figure 23F; cf. "Object-Oriented Contact Patterns"). Stepping and standing on another

animal is distinguished from pawing because the

force of the contact is presumably greater and the

animal establishing contact is usually in a superior

or elevated position. Clasping of another animal's

body occurs in the same manner described for an

inanimate object. As a partner's body is clasped the

hindlegs may also be pumped rapidly back and

forth with the major thrust occurring during the

backstroke. Though both legs may be pumped

concurrently in a reciprocal manner, usually only

one leg pumps at a time while the other is pressed

against the companion's body or held free of contact in a flexed position.

Kicking, the hindleg analog of pawing, consists

of an often rapid extension of the limbs as the animal reclines on its side or back. When contact oc-
50

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

curs it may be forceful and brief or light and of

variable duration.

Trunk

SQUATTING OVER.—Individuals of both sexes are

sometimes seen to straddle the body of another animal with the hindlegs, and then depress the pelvic

region over some part of its body. This may or may

not involve contact; when it does, it is not a forceful pressing movement and there is no indication

that the penis is extended or the perineal gland

everted. Contact of the inside of the upper legs and

genital area seems to be light and intermittent. The

position may be retained without movement or an

animal may creep forward over the other's body

with small steps.

STEPPING OVER.—When one animal steps over the

body of another, it often briefly touches some part

of the latter with the ventrum and or hindlegs. Similar fleeting contact may also occur in jumping over

another animal.

BRUSHING AGAINST.—An animal may briefly brush

any part of its body against that of another animal

in passing.

SOMERSAULTING AGAINST.—Occasionally as two

animals engage in long bouts of contact behavior,

one may place the side of the head on the ground

and perform a lopsided somersault by rolling over

onto the neck, shoulder, and then the side.

Compound Contact Patterns

The behaviors in this category differ from those

described heretofore in being composed of several

independent but concurrent or simultaneous movements. The movements of the two animals may be

alike and reciprocal or different and complementary.

This category is used for practical reasons because

these behaviors are more easily recognized and described at a higher level of organization. The descriptions are simplifications which outline basic

elements of complicated dynamic events.

HEAD-SPARRING.—While frontally oriented, a pair

of animals may mutually swing and toss their heads

in apparent efforts to avoid mouth striking or biting by the companion. The character of the movement resembles that of erratic head movements

except that there is an evident action-reaction relationship between the animals' motions.

GRAPPLING.—Standing bipedally upright, the animals paw at and clasp one another about the

head, neck, shoulders, or chest while head-sparring

or head-darting and biting each other's head, neck,

forelimbs, and shoulders.

WRESTLING.—Animals engaged in this activity

maintain a broad zone of body contact in a head to

head, head to rear, or cross-body orientation. One

or both animals may clasp the other with the foreand/or hindlegs, and paw, pump, bite, or head-spar.

When in body contact, one animal's belly contacts

the other's side, back, or belly.

FIGHTING.—Combat between two animals resembles wrestling but it almost always involves a

cross body or head-to-head orientation and is usually accompanied by screaming, urination, and anal

scent emission of one animal. Wrestling may grade

into fighting, but if fighting movements are more

rapid, body contact is usually briefer, and usually

the initiator repeats the attack. Animals matched

in size, fighting in a head to head orientation, bite

each other primarily on the head, neck, and breast,

and usually manage only to pull out each other's

hair; wounding does not occur. When a considerable

size disparity exists between combatants, the larger

one subdues the smaller by clasping and may bite it

severely.

COPULATION.—The male clasps the female from

behind about the groin and rests his chest and belly

upon her lumbar and sacral region. Intromission is

achieved by the male's pelvic thrusting and the assumption of a lordotic spinal curvature in the female. In one out of three copulations seen, the male

(K) bit the hair on the female's neck (C) during the

terminal seconds and pulled backwards apparently

counterbalancing the precarious position that accompanied final thrusting. In the other two instances (F and I), biting of the female's nape was

not seen.

Companion-oriented Contact Patterns

in Other Genera

The two Civettictis were not seen squatting over,

pumping with the hindfeet, or grappling. Clasping

with either the fore- or hindlimbs is relatively

ineffectual because the wrists and ankles have very

limited powers of rotation. Consequently the African civets are unable to effectively restrain one another by clasping during wrestling or fighting. In

addition, the following contact patterns merit special attention.
NUMBER 239 51

FLEHMEN.—In Civettictis this is an elaborate and

time-consuming pattern usually performed in a

quadrupedal stance during and after sniffing and/or

licking the female's perineal gland, perineal scent,

or urine. The first phase consists of sniffing or licking the scent source; the corners of the mouth may

be slightly retracted, the ears directed laterally, and

the eyes squinted; this is equivalent to Flehmen seen

in Genetta. In the second phase, the head is raised

away from the scent to approximately a horizontal

level and then slowly depressed. If the accompanying facial expression is not assumed during sniffing

or licking, it is assumed during the nonsniffing

phase. Occasionally the eyes are closed and the tip

of the tongue extends over the lower incisors. At

close range a soft, brief, slowly repeated "puff" is

audible, suggesting that the breath is held after

sniffing and expired in small amounts at periodic intervals. The nonsniffing phase is terminated by

licking the nose and lips and sometimes a brief

head shake. This may be followed by another activity or resumption of sniffing.

PERINEAL GLAND SCENT-MARKING.—On one occasion the male Civettictis was seen to evert the

perineal gland, back up, and lightly press it against

the female's neck as she was sniffing the wall of the

cage.

HEAD- AND NECK-WIPING.—While walking past the

female the male Civettictis, on a number of occasions, extended his head or neck toward her and,

with a brief extension movement, wiped some part

of her body.

VOCALIZATION

Table 10 compares the physical characteristics of

the various genet vocalizations described in the following pages.

COUGHING (Muckern: Herter, 1952).—The cough,

a brief sound pulse having a broad energy distribution and a narrow intensity latitude (Figure 27A),

can be mistaken for a sneeze. A true sneeze, however, has a longer duration and is often accompanied

by a characteristic head motion. Coughs may be

phonetically rendered, "uff-viff" (Ducker, 1957), and

are produced singly, in couplets or triplets, or may

be repeated in long series with intervals of various

duration. It could be evoked in all individuals at

the day of birth and continues throughout life.

Production of the sound is preceded by a bumplike

expansion of the throat. Expiration of air through

the glottis and out the nostrils is accompanied by

the disappearance of the bump and a single jerking

compression of the rib cage. Spectrograms were

occasionally obtained from animals O, K, and C

showing a weakly harmonic pulse and a wide noise

band that seem to fit the apparent glottal and nasal

events, (Figure 27A).

HISSING (Fauchen, Explosive Hiss, Spucken).—

This vocalization, phonetically rendered as a noisy

"hcchhh," is another brief duration call having

a broad energy distribution. It is produced by

the rapid expiration of air through the opened

mouth, and the extent of the mouth opening seems

to be correlated with the intensity of the call. During violent hissing the mouth assumes an angle of

about 35°-40° and the jet of air from the glottis

blows saliva out the mouth. Most of the time explosive hissing is accompanied by head-darting with

the glottis opened at or near the end of the extenFIGURE 26.—Facial expression during the panting-hiss in the

genet (animal £; 24 fps; shaded and clear columns = expiration and inspiration respectively, estimated from rib cage

movements; circles = measurements of the jaw-gape angle

in degrees estimated from the film; the curve was fitted by

eye).
52 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
NUMBER 239 53

sion phase. Eye closure may or may not coincide

with the production of this sound but often occurs

if the mouth contacts the recipient.

PANTING HISS.—This call shares many of the physical characteristics of the explosive hiss, but its

temporal organization is a function of accelerated

respiration (Figure 26). The expiration phase of

this sound produces a broad frequency range with

bands of energy concentration resembling dissipated

harmonics, while inspiration produces low frequencies with lower intensity (Figure 27D). The

facial expression of this call is characterized by

retracted lips and a jaw-gape that varies with respiratory phase. In highly aroused animals the position

of the vibrissae during the production of the panting hiss changes the contour of the muzzle (Figure

26); thus, an intense form of the call has an expiration phase accompanied by a widening of the

jaws and an extension of the vibrissae. The retraction of the lips exposes only the lower cheek teeth,

canines, and incisors. The call was first observed at

the age of two days (Fr), five days (M and E), 13

days (O, I, and C), and 16 days (K). Both forms of

hissing may occur with growling and screaming.

WHINING (Nestgezwitscher: Goethe, 1952;

Quicken: Diicker, 1957).—This vocalization varies

widely within and between individuals in harmonics, intensity, and duration (Figures 28A-C). The

leading and/or trailing segment of a whine may be

frequency modulated. Harmonic form often varies

between bouts, and is probably a function of the

animal's activity and general muscle tonus. Calls of

moderate duration have the largest number of harmonics while very brief and very long calls have

fewer harmonics. Whining is phonetically rendered

"wee" or "wew," and is first produced within minFIGURE 27.—Vocalizations of the genet, African civet (Civettictis), and the lesser oriental civet (Viverricula): A, sequence

of three coughs and a whine in Genetta (O); B, series of

eight coughs in Viverricula at the age of 18 days; c, sequence

of calls in the female Civettictis: snort (0.2 sec), growl (0.6-

1.0 sec), and two screams (1.0-1.7, 1.9-2.4 sec); D, panting

hisses in Genetta (M), high frequency hisses occur during

expiration, low frequency hisses during inspiration (six respiratory cycles are displayed); E, purring in Genetta (Fr), the

breaks at 0.8 and 1.7 sec correspond to changes in the flow

of glottal air; F, a growl grading into a scream and returning

to a growl in Genetta (O), from 0.7 to 1.1 sec the scream is

marked by the appearance of a 0.5 kHz frequency band; c,

a growl occurring during a single protracted expiration in

Genetta (M).

utes after birth; it acquires a lower frequency as the

animal matures, and is rarely produced after the

animal becomes independent. Whining is almost

always repetitive and performed with an open

mouth, the extent of the gape varying with the

intensity of the call.

SCREAMING (Schreien).—The scream is essentially

a very high intensity whine but having a greater

number of overtones (Figure 28c). The distribution

of the overtones suggests that two or more membrane surfaces are vibrating at different rates.

Though the call is produced in situations of high

autonomic arousal, the range of intensity subjectively seems to be moderate. Softer calls have fewer

harmonics and, consequently, a narrower range of

energy distribution (Figure 28E, left). The sound

was infrequently heard, but it could sometimes be

evoked by rough handling in animals that had not

yet been weaned. Young animals seem to be unable

to respond to pain or trauma with a sound more

forceful than a loud whine. An open mouth, extended mystacial vibrissae, retracted lips, and dilated pupils compose the facial expression that

accompanies this call. Very high intensity screams

display degeneration of harmonic structure and

broad noise distribution (Figure 28E, right). In

such cases screams appear to grade into noisy explosive hisses.

PURRING (Schnurren).—Purring is composed of

continuous and rapid volleys of low intensity clicks

produced during both inspiration and expiration

(Figure 27E). Usually clusters of sound pulses with

similar interval length are separated from other

such clusters by longer intervals; these breaks in the

sound undoubtedly correspond to the change from

inspiration to expiration. Purring occurs with the

mouth closed or while the animal sucks milk; the

eyes may or may not be closed. It was first heard at

various ages: 2 days (Fr), 5 days (M, E), 13 days

(O, I, C), and 16 days (K). The sound is very quiet

during the first week; this probably explains why it

was not detected in my first animals. Some genets

continued to purr until they were six months old,

but these were raised singly and were exceptionally

tame. As noted by Diicker (1957), purring probably

does not occur in adults.

GROWLING (Knurren).—Growling is a noisy, usually continuous, moderate intensity vocalization

with a repetitive pulse. As in purring, the rhythm of

growling conforms to the period of the respiratory
54 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 10.—Characteristics of genet vocalizations (frequency in kHz, duration in seconds,

duration range in parentheses)

Type and number of

vocalizations

Cough (0)

(K)

Explosive

28

12

hiss (0) 7

Panting hiss (M) 12

Whine (0)

Scream (0

Purr (K)

(F)

Growl (M)

(0)

94

9

6

6

6

8

Frequency Range

Total

0.85-8.0

0.85->8.0

0.85-7.5

1.4^>8.0

0.5--7.0

0.85->8.0

0.35--1.5

Fundamental

Dominant frequency

0.85- 7.0

0.35->8.0

0.35-4.0 -1.0-2.0

1.4--6.0 -1.4

0.5--4.0 ~0.7

0.35-~3.0

0.85--1.5

Duration

Smallest

discrete elements

< 0.07

--

x=0.18

(0.11-0.27)

--

—

<0.007

<0.007

Call

< 0.07

x=0.13

(0.07-0.27)

variable

>1.0

x=Q.76

(0.11-3.59)

x=0.86

(0.42-1.5)

variable

>1.0

variable

>1.0

cycle, but the sound is only produced during expiration. The duration of each uninterrupted period of sound seems to be inversely related to the

level of the animal's excitation. Periods of growling

in a moderately excited animal may last 10 seconds

between breaths, while a greatly agitated animal

growls for about two seconds before inhaling. Intensity of the sound is similarly related to period.

Growling is often, but not always, accompanied by

an alert facial contour; the ears are directed anteriorly, the pupils dilated, and the mystacial vibrissae extended. It is often associated with hissing and

may grade into a scream (Figure 27F). Though first

heard at the age of 80 days (O), it can probably

be evoked earlier.

VOCALIZATION IN OTHER GENERA

The occurrence of call types within selected viverrid genera is summarized in Table 11.

COUGHING.—Unlike the coughing of Genetta, the

call in Viverricida malaccensis 7 has a clicking quality and is produced in bouts composed of several

cough elements (Figure 27B). Of seven recorded

' A recording of coughing and whining in an 18-day-old

male born at the Chicago Zoological Park, Brookfield, Illinois,

was kindly made available to me by Dr. Robert Hoi with.

coughing bouts, two were composed of six and two

of seven elements, and three bouts contained eight

elements. The intervals separating the cough elements varied from 0.03-10.46 sec. The majority of

intervals (79%) fell within the range of 0.06-10.12

sec. The coughing vocalization in Civettictis civetta,

like that of Genetta, is the most common adult call.

It is emitted with the mouth closed and is always a

rapid series of sound bursts composed of from 2 to

14 elements (Figure 29). The sound is phonetically

similar to, but deeper than coughing in Genetta.

Coughing in the fanaloka, Fossa fossa, has a distinctive quality rendered phonetically as "chuck." The

intervals between coughs are variable; while the

sound is most commonly repeated in a rapid series

as with Civettictis, is is also emitted singly at longer

sporadic intervals.

HISSING (SNORTING).—Hissing or snorting was

heard in all the observed genera with the exception

of Viverricida. Diicker (1957), however, recorded

"spitting" (Spucken) in this species (Table 11).

Snorting constitutes a specialized variant of the hiss

in which the dominant lower frequencies probably

result from vibration of the soft palate. Unlike hissing, however, snorting always has a loud, brief, and

explosive character. As infants are often unable to

produce the most forceful of adult vocalizations, it
NUMBER 239

55

FIGURE 28.—Vocalizations of the genet and the lesser oriental civet (Viverricula): \~c, six variations in the whining call of genet K. (not a series); D, four examples of whining in Viverricula

(not a series); E, three examples of screaming in the genet O (not a series).
56 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

IJ

U14

LL

OlO

LJJ ^

cQ 6

i 2

. 9

•

i

N=67

I x=6.12

Th n

, d N=53

p

-| x=4.96

rh

2 6 10 14 2 6 10

NUMBER OF ELEMENTS PER CALL

FIGURE 29.—Variation in the number of elements composing

coughing series in the male and female African civet (Civettictis).

is possible that hissing may replace snorting in the

early life of Civettictis (Figure 27c; Table 11).

Though snorting or hissing may be associated with

head-darting in all the observed species, limited observations on the male Paguma revealed that snorting was always associated with a modified method of

head-darting. The snort was produced as the animal

extended and raised its neck while swinging the

head upwards. At times the animal would quickly

rise up off its forefeet while snorting.

WHINING (GROANING, BLEATING) AND SCREAMING.

—Four Viverricula whining calls from a recording

of 21 are reproduced in Figure 28D. In form these

calls resemble those of Genetta except that the harmonics are more widely spaced and there is relatively little variation in call duration. The one kHz

TABLE 11.—Distribution of vocalizations in social situations for selected

viverrid genera (+ = present, — = absent, +? or — ? = probably present

or absent but substantiation needed, blank = uncertain; calls divided into 6

major families on basis of overall physical similarity)

Vocalization

Viverrinae Paradoxurinae Hemigalinae

o

fc.

s-

>

>

I/I

•t—

+->

o

+J

+ J

0)

>

<_J

CO

c

•9—

-o

c

<o

z

(/)

s-

^

o

T 3

ro

S-

<o

o.

o

to

•r—

It)

B

3

CT

(O

Q_

ra

1/)

I/I

o

u.

Q} -r—

Cough

Hiss

Snort

Whine

Tremelo groan.

Bleat

Scream

Purr

Growl

Hum

Hoot

Neigh

Summary

+ + +

+ +* +1

+ +? +

+

+ +* +

* Described in literature
NUMBER 239 57

kHzO

FIGURE 30.—Vocalizations of the fanaloka (Fossa) and the African palm civet (Nandinia): A-B,

four examples of the tremelo groan in Fossa (note variation in frequency and temporal structure); c, four examples of screaming calls in Fossa (note variation in harmonic detail); D, four

examples of bleating in Nandinia.
58

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

A

B

C

.5

FIGURE 31.—Growling and humming vocalizations of the genet, the African civet (Civettictis),

the Formosan masked civet (Paguma), and the fanaloka (Fossa): A, humming in Paguma; B,

growling in a juvenile Genetta (O); c, growling in an adult Civettictis (female); D, growling in

a juvenile Fossa.

difference in the fundamental frequencies between

the fanaloka and genet can be attributed to age difference. Whining was not heard in Civettictis but a

recording of the female's scream during a fight is

presented (Figure 27c). This sonogram is representative of six others. All are characterized by an

aharmonic broad frequency scattering of noise. For

Nandinia Diicker (1971) described a loud, sustained

"Fiepen" apparently heard only in young animals.

It was not detected in my young male when he was

received at the age of about two months. Bleating

(Figure 30D), rendered phonetically as "Kraa," refers

to a call that may be a variant of whining, though

Diicker (1971) recognized them as distinct. It is produced with the mouth open and uttered softly but

spontaneously by a motionless or slowly moving
NUMBER 239 59

solitary animal. Diicker noted that a louder version

occurred as a response to "unexpected, sudden contact of the body" (1971:81). Though the male and

female Nandinia fought on several occasions, a

screaming vocalization was not heard.

The tremelo groan and scream in Fossa fossa

usually differ in their physical properties, but both

are uttered with an open mouth and sound like

extremes of a graded continuum (Figure 30A-C).

Though the groan usually differs from the scream

by a more complicated and varied sound structure,

groans can have the discrete harmonic organization

that characterizes screams devoid of vocal noise

(Figure 30: compare A, tracing 1 with c, tracing 3).

Most sonograms of groaning consist of unbroken

series of tightly spaced pulses having broad zones of

frequency emphasis (Figure 30A, tracings 2 and 3, B).

These pulses may resolve themselves into deep narrow chevrons (Figure 30A, tracing 2) or thin, broken,

superimposed horizontal bands (Figure 30B). The

scream of Fossa sounds like that of Genetta: the

fundamental frequency lies in the vicinity of one

kHz and the harmonics are tightly packed (Figure

30c, tracings 1-3). Harmonic structure can also be

completely disrupted (Figure 30c, tracing 4).

PURRING.—This call was only heard in the male

Nandinia at the age of 2-3 months. Unlike Diicker's

(1957) animals it was not heard in the two adults.

GROWLING AND HUMMING.—Figure 31B-D presents growling vocalizations in Genetta, Civettictis,

and Fossa. Low frequency noise and a repetitive

pulse structure are common properties of growling

in all three species. The trace of higher frequency

pulses in Fossa (Figure 3ID) and the predominance

of these elements in Genetta (Figure 31B) are reflections of the young age at which both animals were

recorded. It can be seen that the humming vocalization of Paguma (Figure 31A) has a fine grain

pulse structure similar to the growl of the young

genet. This common physical property probably accounts for the relatively smooth sound quality of the

growl of young animals and the humming call of

Paguma and Nandinia (Table 11). Humming, however, is frequency modulated and has a lower sound

intensity than growling. In both Paguma and Nandinia it usually begins at a low frequency (<1

kHz) and then rises slowly and steadily to a higher

pitch. As no recordings of protracted humming are

available, the higher frequency levels of this call are

unknown. Humming may last from several seconds

to at least a half minute. As noted by Diicker (1971)

the call resembles a siren and pairs of animals seem

to hum together. Duetting was also heard in my pair

of Nandinia. In addition, the male was once heard

to "answer" a distant fire engine siren by matching

the frequency change pattern remarkably.

HOOTING AND NEIGHING.—These two distinct calls

are produced with a partly opened mouth and share

the characteristic of high intensity. Hooting was

heard in an old male Nandinia at the National

Zoological Park. Repeated at a rate of about two

calls/sec, it has a relatively high unvarying frequency, and can be imitated with an owl-like "hoohoo-hoo-hoo" sound. Neighing in Paguma, on the

other hand, is composed of a rapid series of brief

high-pitched sounds that gradually rise and fall

within a narrow frequency range; it is best rendered

as "yip-yip-yip-yip." A bout of neighing is uttered

with a single sustained expiration.

Analyses of Interaction

MATERIALS AND METHODS

Data on social interaction in Genetta were collected on a socially stable group of three adults

(Figure 33), and during encounters between familiar

and unfamiliar pairs in a neutral observation cage

(Tables 12, 13).

Interactions of three adult genets (K, C, and I)

within the home cage were recorded for 90 minutes

on 10 alternate evenings. Animals were fed from 2

hrs 15 min to 3 hrs before observation, which with

one exception began at 1900 hours; on the sixth

evening observations began at 2150. Their activities

were dictated into a Uher 1000-L tape recorder

while the writer sat near the end of the cage. After

30 minutes, a large burlap sack was placed on the

floor of the cage. The sack had the effect of a novel

object and induced sniffing, manipulation, and finally intense interaction between the animals. The

sack was removed after 60 minutes. Finally, the recorded notes were transcribed for compilation.

Encounters between familiar and unfamiliar animals were staged in a plywood cage measuring

1.2 x 1.2 x 2.4 m (4 x 4 x 8 ft) with a wire mesh

top and a 1.2 X 2.4 m plate glass front. Animals

were introduced through doors measuring 30 X 30

cm. The cage interior was covered with several coats

of blue-green semi-gloss paint. The floor, walls, and
60

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

glass of the cage were always sponged with soapy

water between series of encounters or after an encounter in which an animal urinated, defecated, or

emitted anal scent. A cardboard box was placed in

each rear corner of the cage and a large forked limb

was placed lengthwise in the center of the floor.

One branch of this limb rested on the top of the

box on the left. Initially a small entrance was made

in each box, but after a pair of animals interacted

out of sight, the fronts were cut away for visibility.

Boxes were replaced after each series of encounters.

Each pair of animals was usually placed together

every other day until a series of 10 successive encounters was completed. To minimize disturbance

during transfer to the encounter cage, a delay of

from 1 hr 15 min to 6 hrs 50 min preceded the introduction of the second animal (Appendix 1). Animals were alternated as the first to be transferred.

They were transferred in cardboard tubes, burlap

sacks, or wooden boxes; these were often explored

and entered shortly after being placed in the home

cage. Animals that did not enter a container were

transferred in their nestboxes. Some animals had

to be captured ahead of time and held in a small

transport cage to reduce excitement before the second introduction could be made. After recording an

encounter, each animal was allowed to enter its own

nestbox and was transferred and fed. A total of 108

encounters was staged, ranging from 11 min 40 sec

to 52 min 50 sec in duration; 84 (77%) of the encounters lasted 30-40 minutes. Observation time for

all encounters amounted to 62 hours, 49 minutes,

and 32 seconds.

Tape recorded observations began as soon as the

second animal entered the cage. The observer sat

within an enclosed canvas blind viewing the animals

through a 30 X 60 cm window ca. 1 m from the

cage. The room was illuminated with white fluorescent lights, but the blind was dark. Most of the

time the animals seemed unaware of my presence.

The targets and duration of contact behavior on

the companion's body were recorded (Figure 32). It

was not always possible to record whether proximate patterns, such as sniffing and head-darting,

involved contact with the companion. Tape recorded accounts were transcribed to Esterline Angus

charts at 15 cm/min (6 in/min) and behavioral

events were numerically coded. A Model F Benson

Lehner Corporation Oscar and Decimal Converter

linked to an IBM Number 26 Printing Card Punch

neck shoulder back rump

head

tailbase

forefoot foreleg hindleg hindfoot

FIGURE 32.—Body topography used in designating the targets

of sniffing and head-darting during encounters between

genets and African civets (Civettictis): the anogenital region

lies beneath the tail and between the hindlegs posterior to

the penis or vagina; the belly encompasses the ventrum

posterior from the chest and anterior of the genitalia.

was used to transpose information on the charts to

punch cards. Data were analyzed with the use of a

Honeywell 2016 Computer and a CDC 6400 Computer.

The two Civettictis were held separately in adjoining cages measuring 1.8 X 1.8 X 3.6 m (6 X 6

X 12 ft). Encounters occurred when a gate was

opened and the male entered the female's cage; the

duration of nine recorded encounters staged over a

14-day period ranged from 33 min 15 sec to 59 min

24 sec (Appendix 2) and totaled 6 hours, 29 min,

and 4 sec. All but the first two encounters occurred

under infrared light. All animals experienced an

artificial light cycle of 10 hrs dark, 14 hrs light with

onset of darkness occurring at 1430 hours. Feeding

took place at ca. 1330 and encounters began from

15 to 60 minutes after dark.

The observer sat unconcealed at the narrow end

of the cage and notes were dictated into a tape recorder. Data tabulations were based on transcriptions of these recordings. Unless otherwise noted,

data were statistically tested with the Wilcoxon

matched-pairs, signed-ranks test (Siegel, 1956).

RESULTS

Effects of the Encounter Introduction Schedule

on Behavior in Genetta

The order of introduction of the two animals in

a series of encounters could affect the conduct of the

participants and the outcome of the encounter.

Five behaviors were selected to evaluate the effects

on animals of the sequence of introduction into

the encounter cage. Approaching and sniffing the

partner were the two behaviors used to measure
NUMBER 239

social initiative; it was hypothesized that having

explored the cage, a resident animal might initiate

more social interactions than the more recently introduced one. Alternatively, an animal might become quiescent after exploring an empty cage and

exhibit behaviors of a restful state when the partner was introduced. Yawning was used as an index

of inactivity. The final two behaviors, sniffing the

cage and climbing upon the cardboard boxes, were

used as measures of exploration and activity. Such

behavior was hypothesized to predominate in the

second animal introduced to the cage.

The frequencies of each behavior were tabulated

for each animal in two categories: introduction into

the encounter cage before or after its partner. These

paired frequencies for each behavior in which the six

animals participated were tested statistically, and

only sniffing of the cage exhibited a significant difference (P — 0.025). This suggests that the introduction of one animal into the encounter cage

considerably before its partner did not determine

the social initiative of the participants.

General Features of Encounters

The types of interaction patterns and their frequency varied greatly between encounters involving

different animals, but within a series of encounters

for a given pair of animals a certain mode of interaction generally prevailed. The general features of

the encounters are outlined below.

COMPANION-ORIENTED CONTACT.—Frequencies of

contact between the three cagemates (K, I, and C)

provide a base for comparison with those in the encounter situation. Social tendencies of these three

animals are illustrated in the sociograms of Figure

33. The relative frequencies of the four classes of

contact (sniffing, head-darting, licking and nibbling,

and fighting) agree with subjective impressions

gained over almost three years of daily observation.

Contact was most frequently exchanged between the

male and females. This is particularly evident in

sniffing and body contacts (Figure 33A,D), but it is

also seen in licking and nibbling (Figure 33c). Most

licking and nibbling was done by the male (K).

Head-darting was most frequently exchanged between K and C. Though C seldom darted at I, I

directed over 75% of her darts at C, making C the

most frequent recipient of this behavior.

Table 12 illustrates that all forms of contact ex61

N=51

FIGURE 33.—Sociograms of selected contact patterns between

three genet cagemates (animals K, I, and C): A, sniffing

(nasal-contacts); B, head-darting; c, licking and nibbling; and

D, body contacts (pawing, clasping, standing on, and wrestling; N = number of times each animal performed a given

type of contact; arrow thickness is proportional to percentage

of occurrence rounded off to nearest 10%; actual percentages

in decimals accompany each arrow).

hibited in the home cage were also displayed in the

encounters. As might be expected, the relative frequencies of contact differed under the two conditions. The level of contact exchange seen in

cagemates occurred in only two of nine pairings

involving unfamiliar animals (Table 12). Protracted

sessions of playful interaction occurred between K

and Mo and between the two immature males (F

and O). Mutual licking and nibbling was exhibited

only by K and Mo, and in the three series of encounters involving familiar animals (C and K, I and

K, I and C). In all other encounters contact between strange animals predominantly involved sniffing and head-darting. Moreover, in almost all cases

these two activities constituted the majority of the

contact exchanged between animals.

Fights occurred in 5 out of 12 series of encounters,

and were initiated by K and Mo. K initiated a fight

with his cagemate C and with the unfamiliar female

Mo. In both instances, the fight was preceded by an

extensive bout of playful interaction involving

wrestling, biting, mouthing, and limb contact.

These activities became progressively more vigorous; the male repeatedly clasped and bit the female

and finally attacked her violently. The female
62

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 12.—Number and percentage (in parentheses) of 5 common contact behaviors as observed in a series of encounters involving 12 paired arrangements of 6 Genetta tigrina study

animals (only pairs 2, 4, and 5 involved animals already familiar to each other)

Anima

1

2

3

4

5

6

7

8

9

10

11

12

F

0

C

K

Mo

K

I

K

C

I

C

Mo

C

F

I

0

c

0

I

Mo

I

F

Mo

F

1 Pair

c/....

0*....

9

</....

9....

<f....

9....

2

?....

?....

9

c/... .

9

d\...

c/....

9

9....

9

<S

* ..

o*....

Totals

Sniffing

401

444

233

161

238

218

160

92

60

58

58

64

79

1

0

43

5

38

8

30

1

2

6

1

2242

(64.8)

(59.8)

(38.7)

(42.8)

(78.3)

(84.5)

(78.4)

(79.3)

(81.1)

(45.3)

(77.3)

(74.4)

(100.0)

(1.7)

(53.7)

(11.9)

(90.5)

(75.0)

(56.5)

Head-darting

31

62

170

42

16

8

24

8

4

49

17

21

0

57

9

30

33

4

4

7

12

2

0

0

610

(5.2)

(8.3)

(28.2)

(11.2)

(5.3)

(3.1)

(11.8)

(6.9)

(5.4)

(38.3)

(22.6.)

(24.4)

(98.3)

(37.5)

(78.6)

(9.5)

(17.5)

(15.4)

Licking and

nibbling

0

0

58

27

8

19

7

14

1

5

0

0

0

0

0

0

0

0

0

0

0

0

0

0

292

(9.6)

(7.2)

(2.6)

(7.4)

(3.4)

(12.1)

(1.4)

(3.9)

(7.4)

Biting and

mounting

106

56

79

71

14

5

6

1

2

9

0

1

0

0

1

7

3

0

0

3

1

0

1

0

366

(17.2)

(7.5)

(13.1)

(18.9)

(4.6)

(1.7)

(2.9)

(0.9)

(2.7)

(7.0)

(1.2)

(8.8)

(7.1)

(7.5)

(9.2)

Limb

contact

80

181

62

75

28

3

7

1

7

7

0

0

0

0

0

0

1

0

0

0

0

0

1

0

458

(12.9)

(24.4)

(10.3)

(19.9)

(9.2)

(3.1)

(3.4)

(0.9)

(9.5)

(5.5)

(2.4)

(11.5)

Total

618

743

602

376

304

253

204

116

74

128

75

86

79

58

10

80

42

42

12

40

14

4

8

1

3968

screamed and attempted to escape for about a minute before the male desisted. Mo initiated three

attacks without prior wrestling.

Adult males must be kept separate to avoid possibly fatal hostility. On occasions when either O or

F were moving freely about the laboratory, K responded to their presence with unmistakable signs

of aggression. When an unrestricted animal approached the cage, K would rush toward it and

follow its movements by running along the inner

boundary of the cage. On two occasions the male's

excited demeanor caused the resident females to

run also. The motion of the two female cagemates

and the outsider appeared to confuse the male who

then chased a cagemate until he caught and attacked her. For this reason, K was never introduced

to either of the other male genets.

NONCONTACT PATTERNS.—While sniffing and headdarting are the most frequent forms of contact,

they usually occupy a relatively small amount of

the time that two animals are together. An animal's

noncontact activities are also a potentially important source of information to the companion. In

particular, the time-space characteristics of a number of patterns have obvious social significance. The

method of data recording, however, did not permit

consistent precision in assessing the duration and

graded character of these patterns, so the following
NUMBER 239 63

TABLE 13.—Number and percentage (in parentheses) of companion-oriented locomotion, stances,

and movements as observed in a series of encounters involving 12 paired arrangements of 6

Genetta tigrina study animals (only pairs 2, 4, and 5 involved animals already familiar to each

other)

1

2

3

4

5

6

7

8

9

10

11

12

Animal

pair

F

0

C

K*

Mo

K*

I

K

C

I

C

M o*

C

F

I

0

C

0

I

M o *

I

F

M o *

F

</"••

9 "

<r •

9 ••

4 '•

9 ••

<f ••

8 . .

9--

9..

9 ..

9..

cf..

*••

9-.

<J..

9.-

9--

9--

<£••

9--

</••

Approaches

Crouch1

1

13

4

10

35

25

0

0

0

0

10

58

6

3

9

33

2

25

11

4

0

6

1

1

(0.5)

(7.9)

(3.9)

(8.6)

(74.5)

(14.8)

--

--

(12.9)

(38.4)

(8.6)

(42.8)

(69.2)

(12.1)

(13.3)

(22.9)

(15.7)

(6.3)

(12~7)

(20.0)

(50.0)

1

13

6

7

11

9

0

0

0

0

2

8

0

2

1

39

5

0

3

1

0

3

0

0

Rush

(0.5)

(7.9)

(5.9)

(6.0)

(23.4)

(5.4)

--

--

(2.6)

(52.9)

(28.6)

(7.7)

(14.3)

(33.3)

(4.3)

(1.6)

(6~4)

--

Total

199

163

101

116

47

168

60

38

28

58

77

151

70

7

13

111

15

109

70

63

2

47

5

2

Departures

total

235

260

137

171

141

123

51

36

50

45

59

93

16

71

3

160

10

78

53

55

31

3

3

1

Deviation

from a

proportion

of 50:502

App.

5.0

3.5

29.2

11.2

17.4

16.2

40.9

45.4

47.9

2.6

45.9

71.4

Dep.

2.5

4.5

3.4

8.6

2.6

11.2

31.6

48.2

38.6

0.9

41.2

25.0

Visual

fixation

19

36

20

35

132

97

5

12

7

10

173

179

64

78

82

221

95

168

147

105

87

49

7

6

Jaw

gaping

0

0

33

3

6

3

0

1

2

0

12

3

0

49

19

1

34

2

58

4

115

0

0

1

Archedback

stance

0

3

0

1

1

2

0

0

0

0

9

0

1

0

2

3

1

0

10

6

9

1

0

0

Tail

flaring

0

0

7

13

5

19

2

1

1

0

22

23

8

1

2

11

8

8

20

27

3

1

1

C

* Encounters in which fighting occurred; asterisk indicates animal that initiated and won fight.

Includes crouched stances.

2

 Calculated by subtracting from 50.0 the lowest percentage an animal contributed to the approach or departure

total for that pair. The value is placed in the row of the low scoring animal. Deviations <_ 5.0 are placed

between rows.

qualitative description will have to suffice as a rudimentary coverage of this aspect of communication.

Before considering some of the details of interaction characterized by minimal contact, the protocol of an encounter will be summarized. This

observation ought not be interpreted as "typical,"

but should illustrate some salient noncontact features. The example is encounter number 5, 1930

hours; the subjects are Mo (female) and I (female)

(Appendix 1).

1. Mo leaves nestbox; I reclining within her own nestbox

and watching Mo; Mo approaches I, naso-nasal contact, turns,

departs, enters nestbox; Mo's tail = y4 expanded, I's tail

= 0.
64 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

2. Mo leaves nestbox, smells cage; I watching Mo; Mo returns to nestbox.

3. Mo leaves nestbox, tail = i/2 expanded, approaches I

to ca. 30 cm, smells cage, departs; approaches I again, looks at

her, moves about cage, tail flares to 2/3, enters nestbox.

4. Mo leaves nestbox, tail slightly <i/i expanded; I still

watching, assumes crouched stance; Mo backs up, turns away,

flares tail to z/s, then to y4, smells cage, sits, tail = i/2; I

still staring in crouched stance; Mo stands and enters nestbox;

I assumes reclining position; Mo yawns and sits down.

5. I leaves nestbox, tail = 0; Mo looks at I, I stops moving,

looks at Mo, returns to nestbox, sits, then lies down.

6. Mo leaves nestbox and smells cage, tail = 0, climbs on

top of I's nestbox, smells it, departs, tail flares to i/£, enters

her own nestbox and sits.

7. I leaves nestbox and smells floor, tail = 0; Mo watches

her; I approaches Mo, sits down and looks at Mo, stands and

assumes a weak arched-back stance with a tangential orientation to Mo, flares tail to yt

, departs, climbs on top of her

own nestbox, smells cage, tail = \/A, steps down from nestbox;

Mo still watching motionlessly; I walks to front center of

cage, looks at Mo, flares tail to 2/s, turns away, returns to top

of nestbox, tail = i/J; Mo still watching; I yawns and steps

to the ground; Mo turns her head slowly while watching

I .. . gaze is fixed on her; I approaches Mo to within 30

cm, smells toward her, turns away, departs, tail flares to y4,

walks to front center and back to nestbox 2 times; Mo staring

intensely; I's tail is i/2 expanded; Mo rushes at I, I departs.

Mo chases her about the cage, bites her, they fight, Mo bites,

chases, bites, grabs her on the back and bites her on the

neck; I escapes and ricochets off the cage walls; more biting;

they stop and both animals assume lateral body orientations

and stare at each other in strongly arched-back stances; tails

are fully expanded; I runs into her nestbox growling, looks

out at Mo who is smelling urine, anal scent, and tufts of

hair on the floor. Mo enters her own nestbox, smells the

inside and then sits, tail = % expanded; I still maintains an

arched-back stance and growls. Termination.

There are several notable features of this encounter: (1) the two animals exhibited unequal

amounts of activity, (2) nestboxes were foci of rest,

(3) the active animal was watched intently, and (4)

body stances and hair postures were sometimes

modified during visual fixation.

Activity of the two animals during encounters

involving minimal contact is usually nonoverlapping and one animal often predominates. This

results when one animal becomes active more frequently or for longer periods of time; when the

companion becomes active it usually elicits activity

in the dominant animal. The companion then usually retreats and avoids concurrent activity. It is

difficult to make general statements regarding noncontact patterns on the basis of frequency alone;

undoubtedly a number of other variables determine

the appearance of these behaviors.

Three patterns of approaching and departing

were observed in the encounters recorded in Table

13. First, the relative numbers of these actions by

each animal may be approximately equal, indicating equal activity. In minimal-contact encounters

(pairs 6-12), the activity of the participants is largely

out of phase, with neither animal (e.g., Mo or I)

predominating. Second, both approaching and departing may be performed primarily by one animal;

in the minimal-contact situation this simply reflects

that one animal dominates activity to the exclusion

of its companion (C and O, I and O, C and Mo).

Third, when approaches and departures are most

frequent in different animals, the animals may be

supplanting each other. In the minimal-contact encounters (C and F, I and F), the inactive animal

almost always departed to curtail contact with the

active companion.

A comparison of approaching frequencies (Table

13) with sniffing frequencies (Table 12) reveals that

in cases where crouching and rushing intimidation

occurred (pairs 1-3), the animal who approached

the most sniffed and head darted the least. In

minimal-contact encounters, with one exception (I

and Mo), the approaching animal sniffed the most.

This feature of minimal-contact encounters appears

real because the frequency disparity between animals was large for both sniffing and approaching

(Tables 12, 13). Among high-contact encounters, the

subequal frequencies of approaching and sniffing

suggest there is no correlation between sniffing and

approaching.

Crouching and rushing, taken together, appear to

be associated with social initiative. Usually animals

that most often crouch and rush also sniff their companions the most (compare Tables 12 and 13). In

the 2 out of 9 applicable cases the deviation from

a 50:50 approach proportion was <5.0% (Table 13).

Because arched-back stances are of long duration, it

is difficult to evaluate their relationship to the other

patterns by considering the frequencies in Table 13.

In any event, animals that maintain protracted

stances crouch and rush less frequently than their

companions, and hence usually display a low social

initiative. Both crouching and the arched-back

stance clearly have an intimidatory function and

give the impression that crouching is offensive and

back-arching defensive.

Though visual fixation and jaw-gaping are also
NUMBER 239 65

duration events, these behaviors are briefer and

more frequent than the arched-back stance, and

therefore more easily related to other patterns. In

7 out of the 10 applicable encounter series, the animal that stared more than its companion also

crouched and rushed more. In instances where this

animal also gaped its jaw more, it had been defeated

in a fight (Mo by K, and Mo by I).

Visual fixation was generally most common

among animals exhibiting minimal contact (Table

13, pairs 6-11) and among animals that fought

(pairs 2 and 5: C and K, Mo and K). The lowest frequencies of visual fixation occurred in encounters

between the three familiar animals, C and I, I

and K. Pair 12 in Table 13 is discounted in this

case because encounters were discontinued after the

fight.

Jaw-gaping was relatively infrequent in familiar

animals that did not fight (I and K, C and I). In the

minimal-contact encounters (Table 13, pairs 6-11),

a marked disparity in jaw-gaping is evident between

the two participants varying in magnitude from 4

(C and Mo) to 14 times (I and Mo). Pair 12 is again

discounted for the reason stated above. Among animals that fought, the loser jaw-gaped the most.

These were also the animals that approached their

companions less frequently (excepting I and Mo).

Flaring of the tail hair occurred in both high- and

minimal-contact encounters (pairs 1-5, 6-12). High

frequencies of tail-flaring were seen in four of five

encounter series where fighting occurred (the exception was the brief discontinued encounter between

Mo and F, pair 12). Tail-flaring was infrequent both

with familiar animals that did not fight (I and K, C

and I) and with the pair that played (F and O). A

comparison of tail-flaring and jaw-gaping (Table 13)

reveals that the animal that jaw-gaped the most usually also tail-flared the most, but the relationship is

not significant (P > 0.05, Appendix 3). Tailflaring often occurred as an animal maintained an

arched-back stance, but this relationship is not

shown in the table.

CONTINGENCIES OF COMMON NONVOCAL INTERACTION PATTERNS IN Genetta.—Table 14 gives a rough

approximation of some common behavioral associations. The table is an abbreviation of the original

matrix, which contains twice as many behavior categories. The consequence of eliminating many of the

fine grained behaviors (e.g., closes eyes, ears back,

mouth open, etc.) is that some common dyads are

obscured. Sniffing and darting is a case in point.

This common dyad is not shown in Table 14 because the intention movements of darting were recorded between the occurrence of the sniff and the

head-dart. Several contact patterns are presented as

having followed depart (e.g., sniff, dart, bite, paw,

and stand-on). This resulted from failure to record

all approaches, particularly during fast action. In

addition, seemingly incongruous sequences like

these occurred when an animal attempted to depart. For example, as an animal turned to depart,

its tail might be sniffed by the companion.

The simplest form of interaction during encounters consisted of movement toward and away

from the companion. Companion-oriented movement was frequently interposed with sniffing by one

or both animals. Sniffing and head-darting often

occurred in minimal- and maximal-contact encounters and constituted the most common elements of

social exchange. For convenience the association

between these two patterns will hereafter be referred to as the sniff-dart axis.

The sniff-dart axis and multiple contact exchanges represent the ends of a complexity continuum involving contact. Though it is possible for

any two patterns to occur either singly or in repeating and alternating series, sniffing and darting

were by far the most common elements in dyadic

interaction. Other types of contact (e.g., pawing and

biting) had no dyadic integrity.

Biting, pawing, standing-on, and erratic head

movements are relatively infrequent behaviors during most encounters, and their contingencies are by

no means fixed. Tail-flaring is almost always seen in

minimal-contact situations in association with movement away from the companion (Table 13). The

arched-back stance tends to repeat itself as an initial

response to the visual fixation and approach of another animal. It can be seen that noncontact patterns such as tail-flaring, the arched-back stance, and

sometimes erratic head movements are associated

with movement toward or away from the companion, but the first two patterns are at times

precipitated by fighting. Erratic head movements

were subequally contingent upon contact, sniffing,

and companion-oriented locomotion, indicating

that the pattern was performed during, as well as

between, episodes of contact.

CONTINGENCIES OF COMMON, NONVOCAL INTERACTION PATTERNS IN Civettictis.—The two Civettictis
66 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 14.—Contingency matrix of interaction patterns in Genetta tigrina based on encounters

of 12 paired arrangements of 6 study animals; matrix includes behavior patterns from sequences

enacted by one animal and/or between two animals (App = walking approach, Appc =

crouched approach or stalk, Appr = running approach, SO = stand on or clasp with forelimbs, EHM = erratic head movements, TF = tail flare, ABS = arched-back stance)

Fol 1 ows ^""~-\

App

Appc

Appr

Sniff

Dart

Bite

Paw

SO

EHM

Depart

Follow

TF

ABS

TOTALS

App

19

18

12

45

2

1

3

84

6

9

1

200

Appc

28

00

3

5

1

45

Appr

6

2

1

9

Sniff

241

10

323

4

1

1

3

6

30

18

6

643

Dart

4

1

5

36

1

6

1

54

Bite

1

4

3

3

3

2

1

17

Paw

3

4

1

1

9

SO

2

1

2

8

3

1

17

EHM

8

1

6

2

1

4

22

Depart

97

12

2

177

8

5

2

2

7

26

3

21

1

363

Follow

8

1

17

3

1

1

9

1

7

48

TF

4

2

2

1

24

7

6

46

ABS

2

1

5

8

TOTALS

423

54

26

595

55

14

5

9

19

181

34

53

13

1481

exhibited behavioral contingencies similar to Genetta but different by the incorporation of several

other patterns. As with genets, departure frequently

terminated sniffing, and sniffing and head-darting

were both repeated more often than they were followed by other patterns (Table 15). Aside from

such repetitions, sniffing was most frequently

followed by head-darting. Unlike the genets,

however, the recipient of head-darting more

commonly reclined than departed from the companion's vicinity. Under these circumstances, recumbancy characteristically involved a lateral or

tangential body orientation to the companion. This

kind of body orientation was also assumed by standing animals; the pattern was most commonly a

response to the head-darting or frontal orientation

(turn towards) of a proximate companion. Sparring

and clasping were also common antecedents of reclining, but only in the former was reclining associated with a lateral body orientation. It was evident

in these two animals that the assumption of either a

standing or reclining lateral orientation often precipitated further darting. After the darting animal

departed or desisted, the reclining one would stand

and move away. Approaching, turning away, departing, and following bear a basic similarity to the

same patterns in the genet. Erratic head movements

were not commonly seen, but occurred most often

before an animal moved away from its companion.

In both Genetta and the Civettictis sniffing and
NUMBER 239 67

TABLE 15.—Contingency matrix of interaction patterns in Civettictis civetta based on nine encounters of 2 opposite sex study animals (EHM = erratic head movements, LO — lateral or

tangential body orientation, TT = turn towards, TA = turn away, boxed figures = values that

account for 20% or more of column totals)

^\Precedes

Follows ^ \

App

Sniff

Dart

Spar

B1te

Clasp

EHM

Recline

Depart

Follow

Stand

LO

TT

TA

TOTALS

App Sniff

125 1 108 |

47

6

7

23

5

26

17

2

4

13

17

3

295

27

7

2

7

6

24

96

26

6

37

10

356

Dart

14

140

16

4

7

73

48

19

32

10

15

383

Spar

1

12

5

1

19

21

8

4

1

72

Bite

1

4

2

1

2

2

2

13

2

3

2

34

Clasp

2

4

11

6

1

3

21

8

2

1

1

60

EHM

1

1

1

1

3

2

4

10

3

2

1

29

Reeli ne

4

40

47

8

4

3

5

11

26

I

42 1

15

3

208

Depart

1 32 1

11

13

2

97

Follow

8

4

1

3

1

6

7

7

37

Stand

4

4

13

4

5

4

3

7

65

2

2

10

123

L0

3

21

5

4

18

1

3

3

58

TT

3

13

40

13

3

16

12

3

14

1

5

123

TA

5

4

6

12

7

2

2

38

TOTALS

43

325

364

73

36

54

30

235

353

41

132

71

110

46

1913

head-darting were more commonly contingent upon

themselves than upon any other single pattern of

contact. At this level, interaction in Civettictis appeared to be more complicated because of the assumption of postural modifications that permitted

further head-darting; proximal interaction was usually terminated when the head-darting animal departed. In Genetta head-darting and sniffing were

terminated when either animal departed. Lateral

body orientation, assumed during back-arching, was

characteristically displayed at a distance rather than

in the companion's proximity. In addition, Genetta

modified its movement toward and away from the

companion with postural adjustments such as

crouching and tail-flaring.

CONTINGENCIES OF VOCALIZATIONS IN Genetta.—

As mentioned previously, coughing in Genetta was

usually produced in bouts with intervals ranging

from one to several seconds. It is evident in Table

16 that more than half of all coughs were preceded

or followed by vocalizations, the majority of which

were coughs. Locomotory transitions, and movement to and from the companion accounted for

nearly 30% of all contingent behaviors; about 15%

of the calls occurred either before or after the animal sniffed the cage or its companion. Although
68 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

sometimes a response to another animal's contact,

an outstanding feature of this vocalization was that

it commonly occurred during transitions in gaits,

stances, reclining positions, and sniffing. Once an

animal produced the sound, it was likely to do so

again before a change occurred in its ongoing behavior.

Hissing also occurred most frequently in series;

of the 110 times it was contingent upon vocalizations (Table 16), only 10% involved another call

type, i.e., growling. There was a fourfold difference

in the percent of the time that hissing was preceded

and followed by companion-oriented locomotion;

this is because the call was usually produced as a

response to a companion's approach. Darting was almost always performed by the vocalizing animal and

was twice as common after the call as before. About

25% of the time either visual fixation or jaw-gaping

followed the delivery of a hiss; both of these acts

were displayed predominantly by the caller. It is

clear that hissing occurred primarily during proximal interaction in which contact was minimal.

In the encounter situation, growling was only

exhibited by animals that had been defeated in

fights. Following defeat, animals often growled

merely at the sight of the companion indulging in

"neutral" activities, such as self- directed licking and

nibbling, cage sniffing, or changing body position.

TABLE 16.—Number and percentage (in parentheses) of selected behavioral contingencies

with vocalizations in Genetta tigrina

Contingent

behaviors

Sniff cagel

Locomotion2

Companionoriented

movements

Arched-back

stance

Visual

fixation

Jaw-gaping

Head-darting...

Other

contacts

Vocalizations..

Stretching,

shaking and

self-oriented

contact

TOTALS3

Cough

Preceded

by

45

97

69

—

--

—

49

6

316

25

607

(7.4)

(15.9)

(11.4)

--

--

-

(a D

(0.9)

(52.1)

(4.1)

Followed

by

56

96

73

--

--

—

45

6

316

9

606

(9.2)

(15.8)

(12.8)

--

—

--

(7 A)

(0.9)

(52.1)

(1.5)

Hiss

Preceded

by

7

--

33

2

7

14

I

25

1

58

148

(4.7)

—

(22.3)

(1.4)

(4.7)

(9.5)

(0.7)

(16.9)

(0.7)

(39.2)

—

Followed

by

0

—

8

1

16

20

45

1

52

143

—

—

(5.6)

(0.7)

(11.2)

(13.9)

(31.5)

(0.7)

(36.4)

-

Growl

Preceded

by

2

1 "3

11

1

5

5

11

2

4

6

60

(3.3)

(21.6)

(18.3)

(1.6)

(8.3)

(8.3)

(18.3)

(3.3)

(6.6)

(10.0)

Followed

by

3

f.

2

1

8

12

14

2

__

3

5

56

(5.4)

(10.7)

(3.6)

(1.7)

(14.3)

(21.4)

(25.0)

(3.6)

(5.4)

(8.9)

Scream

Preceded

by

—

--

14

—

—

3

18

3

38

—

--

(36.8)

—

__

—

(7.9)

(47.4)

(7.9)

—

Followed

by

—

--

3

—

—

2

25

3

33

—

--

(9.1)

—

-

(6.1)

(75.8)

(9.1)

1 Includes licking and nibbling the cage or scent marks.

2

 Including transitions from stances to locomotion.

3 Estimates of degree of tail-hair erection were occasionally contingent upon vocalizations The

figures results in unequal totals.

exclusion of these
NUMBER 239 69

Defeated animals also growled as they paced

in a stereotyped manner in front of the cage.

Growling occurred before as well as after animals

visually fixated their companions, or assumed an

arched-back stance. These three behaviors accounted

for 38% of all antecedent acts and 46% of the succedent acts (Table 16); hissing was the only vocal

pattern contingent on this call.

Screaming was the least frequently heard call during encounters. It occurred before or after the reception of head-darts, but was most frequently

emitted as animals engaged in wrestling, fighting,

biting, and clasping. Contact patterns accounted for

9

BEFORE

9 a

AFTER

B

9 d

AFTER

9 cf

BOTH

n _ n n

r

a

dartspar h 5 2

change body pos. g 1

recline f 9 3 4

approach e

depart d 4 1 2 4

latorient c 2 1

frontal orient b 8 1 1

no change a 13 3 7 3 2 2 2

41 5 13 3 14 2 2 1

6 0 -

5 0 -

30

20

10-Lm.

i I

50%-75% of all antecedent and succedent behaviors. Approaching was the only form of companionoriented locomotion that animals responded to by

screaming. Screaming then differs from growling

and hissing by its association with extensive body

contact. The contingencies (Table 16) do not indicate that pain inflicted during biting is the most

important stimulus evoking this call. Associated

patterns such as clasping and wrestling were never

seen to elicit the call by themselves. It appears that

once an animal received a painful bite, however,

more generalized patterns such as approaching became effective stimuli.

CONTINGENCIES OF VOCALIZATIONS IN Civettictis.—

As in Genetta, Figure 34A shows that sniffing the

cage and general body movement constituted the

vast majority of activities associated with coughing.

It can be further seen that in each animal olfactory

and (less commonly) gustatory sampling of the cage

was quite often an antecedent to vocalization. Less

than 20% of all coughing-contingent behavior was

nasal contacts. In both animals less than 95% of all

nasal contacts was contingent upon coughing. Locomotory transitions were relatively more frequent

succedents than antecedents.

More than 70% of the occasions after a call was

FIGURE 34.—Behavioral contingencies of vocalizations in the

African civet (Civettictis):

A, Caller's behavior before, after, and both before and after

coughing (the segments of each bar in the graphs represent

different behavior patterns: bottom segment = sniffing, licking, and nibbling of the cage (or scent marks); middle segment = sniffing of the companion; top segment = transitions

in locomotion and stances; difference between bar and 100%

indicates percent of calls with no overt contingent behavior;

numerals above each bar represent the number of calls from

which percentages were calculated).

B, Companion's behavior before, after, and both before and

after the caller's coughs (bottom shaded segment •» nasal

contacts; lower clear segment = all other companion-oriented

contacts (bite, dart, paw, stand on, etc.); middle shaded segment = approach, turn toward, look at; upper dear segment = depart, turn away, look away; top shaded segment

= coughing).

c, Behavior patterns antecedent and succedent to growling

by the female (the bottom graph displays patterns of the

male—white bare—and female—black ban—that were antecedent to growling by the female; the succedent events in the

male are enumerated above each bar; upper graph presents

the relative proportions to those succedent events expressed

as percentages of the total number of times the female

growled, the lower case letters correspond to the behavior

patterns listed at left).
70

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

given there was no overt change recorded in the

companion's behavior (Figure 34B). Antecedent activities of the companion may have evoked the call

on some occasions. It is evident that contact by the

female was the preponderant antecedent to the

male's coughing; and that moving or looking toward

or away from the partner was the chief activity of

the male preceding the female's coughing. Differences between animals were also exhibited in types

of response. In response to the male's coughs, the female sniffed the male 3%, approached 9%, departed

2% of the time, and never coughed back. After the

female called, the male contacted her 15% of the

time, approached 5%, and coughed 7% of the time.

The contingencies of growling differ considerably

from coughing. Of 85 growls by the female, 80%

were apparent responses to actions of the male

(Figure 34c, bottom). Female-oriented movements

and nasal contacts accounted for 81% of all the

male's actions that evoked the female's growling;

she also growled when the male simply moved.

When there was no change in the male's behavior,

the female commonly growled when she moved in

his direction.

It is evident that nearly 40% of the time after

the female growled there was no change in the

male's conduct (Figure 34c, top). The most common

form of response consisted of position or orientation

changes (e.g., reclining, lateral and frontal orientation, and departure). Growling led to contact in the

form of sparring only 10% of the time.

SENSORY CHARACTERISTICS OF SOME NONVOCAL BEHAVIORS.—As a nocturnal forest species having a

retina composed solely of rods (Diicker, 1957),

Genetta distinguishes only degrees of brightness and

probably cannot discern fine visual detail. In view

of these limitations, it is important to consider the

visual characteristics of noncontact patterns and

how they might be perceived by conspecifics. The

patterns discussed below can be recognized only

visually. Noncontact patterns involving gaits (approach, departure) may also have important visual

components, but concurrent environmental sounds

produced by footfalls also provide information concerning proximity, and direction and rate of movement.

Staring is a common activity during encounters

between strange animals. If visual fixation provides

information only to the animal engaging in it, there

would not be selection for behaviors or structures

that emphasize this activity to the companion. On

the contrary, the patterning of light-colored fields of

facial hair provides two characteristics hypothesized

as facilitating the companion's recognition of the

head and its orientation in dim lighting. (1) The

markings present a distinct visual configuration not

present elsewhere on the body, and (2) angular deflection of the head from a frontal position is precisely indicated by relative changes in the number

and surface area of the markings. Figure 35 shows the

appearance of different head positions and provides

measures of the amount of white facial hair exposed

through a 180° horizontal arc. It is evident that

disruption of pattern symmetry and disparity in

spot size are two manifestations of only a 30°

deflection from a frontal head position. These

changes occur with about a 20% decrease of the

area of white hair exposed during frontal orientation. As the eye has relatively little mobility (see

"Primary Senses"), visual fixation of movement is

achieved primarily by adjustments of head position.

A moving animal being stared at therefore probably receives a constant facial image in which four

symmetrical light markings are prominent. Ironically, a similar effect may also be produced by an

animal with closed eyes if the head is directed

toward the companion.

Staring accompanies crouched and arched-back

stances, but the visual characteristics of these patterns are distinct. The body axis is frontally

oriented during crouching, exposing a relatively

small body area to the companion. The tail is concealed or minimally visible and the abovementioned facial characters predominate. On the

other hand, a lateral or tangential position maximizes body exposure in the arched-back stance. The

light-colored hair on face and tail delimits the

animal's dimensions. Differences between these two

patterns may be considered antithetic (Darwin,

1872) and involve both quantitative (head elevation,

degree of body exposure) and qualitative differences

(visibility or nonvisibility of tail).

The rapid flaring of hair on the tail produces

several changes that may be perceptible to a companion. Because the bases of light hairs are lighter

than the tips, erection seems to increase the contrast between the dark and light colored bands.

Pilo-erection also noticeably enhances the demarcation between bands; in the relaxed state the overlapping of light and dark hair at band boundaries
NUMBER 239 71

FIGURE 35.—Appearance of different head positions in the

genet and variation in the relative proportions of the white

facial markings through a 180° horizontal arc from a frontal

position; (gray horizon delimits the area of exposed white

hair as a percentage of the amount visible at 0°; approximate

percentages of the muzzle and eyespot markings are indicated by the solid and dotted lines respectively, together

they equal the percentage expressed by the gray horizon; calculations are based on planimeter measurements from enlarged photos of the horizontally resting head of a dead

genet).

produces a more graded effect. In all probability

the visual pattern that is produced—a rapid expansion of a series of light-colored bars—draws attention to the tail and the animal.

The sensory properties of other behavior patterns

are more difficult to enumerate. The properties

actually transmitted are not necessarily those perceived by the observer and probably vary under

different environmental and social circumstances.

Head-darting, for example, involves several levels

of motor integration. In its simplest form it may be

visual or tactile. If the companion is watching the

darting animal, the signal can exhibit visual, as well

as tactile and auditory properties. The sensory manifestations of the transmitted signal, therefore, depend upon the receiver's body orientation and

attentiveness. The redundant character of such a

pattern probably compensates for interference of

one or more sensory channels. Sudden high stimulus

contrast characterizes all the sensory properties of

this pattern.

It appears that the most obvious properties of

certain patterns may not always embody the salient

stimuli. Though erratic head movements are usually

conspicuous in a lighted room, they are commonly

performed in circumstances where visual detection

is not possible. The irregular and noisy sequence

of footfalls is an outstanding feature of this movement detectable irrespective of orientation to the

other animal. In a sense the quality of this sound

mimics the visual character of the movement. When

the movement is performed during following, the

head and neck glance and brush against the rump,

hindlegs, and tail of the anterior animal.

BODY TARGETS OF SNIFFING AND HEAD-DARTING IN

Genetta.—Figure 36 reveals a bimodal distribution

of sniffing frequencies to the fore and hindbody.

While the head, nose, and neck were the most commonly sniffed forebody targets, the rump, anogenital

region, and tail (including the tailbase) received

45.2% of all nasal contacts, the tail and tailbase accounting for 29%. These common anterior and

posterior targets accounted for exactly two-thirds of

all companion-oriented sniffing. The side was the

remaining most commonly smelled anterior target

region; it also constitutes a relatively large surface

area. Forty-six percent of all sniffs were directed to

targets between the head and side, while the remaining parts of the body, composing slightly less

than half the available body surface, received 54%

of the total number of sniffs.

The ratio of the frequency to the total duration

of sniffling for any given target varied considerably

(Figure 36). Ratios of <1:1.5 were obtained for all

targets anterior to and including the side. On the

hindbody only the legs and feet had similar ratios.

Almost all other posterior body targets had fre-
72 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

172 167 196

54 298 300 312

HEAD-DARTING

89 -

3 2 9 4 1

66 70

96 88 1 J

i m^ 4

tailbase tail hindtoot _

hindlegV un-spec

FIGURE 36.—Body targets of head-darting and sniffing in the genet (numerals above bars in headdarting graph and top numerals above bars in sniffing graph = number of observations for that

target; bottom numerals above bars in sniffing graph = total duration in seconds of sniffing

directed to each target; solid bars = frequency percentages based on total number of all observations for specified and unspecified targets; shaded bars = duration percentages based on

total duration of sniffing directed to all specified and unspecified targets).

quencyrduration ratios of 1:1.5-1.9. The spinal crest

had the exceptional ratio of 1:3.75. This clearly

demonstrates that duration and frequency of contact

are not equally proportional for all targets. Frequently sniffed targets such as the head were sniffed

briefly (frequency:duration = 1:1.3), while relatively long periods were devoted to infrequently

sniffed targets such as the spinal crest (frequency:

duration = 1:3.75). Spinal crest sniffing exhibited

a characteristic pattern: the nose (probably guided

by the contour of the underlying vertebrae) was

rubbed anteriorly within the confines of the black

hair of the crest.

The vast majority of the time the companion's

head was the head-darting target; the neck, forelegs, and tail were struck at less frequently. Other

body areas were infrequent targets (Figure 36).

BODY TARGETS IN Civettictis.—An interesting

comparison with Genetta is presented by this

species. While the head, neck, and sides were the

most commonly sniffed anterior regions, in the pair

of Civettictis the majority of sniffing was directed to

posterior body targets. The female directed 60% of

all nasal contacts to the male's hindquarters, while

the male addressed 65% to this area of the female's

body (Figure 37).

In a number of instances it was possible to determine the precise orientation of sniffs to specific body

FIGURE 37.—Body targets of head-darting and sniffing in the

female and male of Civettictis (each category exclusive of

others): A, general body targets of head-darting and sniffing

in the male and female; B, specific targets of head-darting

and sniffing on the head of the male and female; c, specific

targets of sniffing on the anogenital region of the male and

female (female not shown here to conserve space; because body

targets were not always noted specifically during an encounter,

general targets such as head and anogenital region have been

duplicated in B and c, e.g., the undertail category of c was included as part of the anogenital category in A; numerals accompanying each segment of the graph represent the number

of observed instances from which the percentages are calculated).
NUMBER 239 73

4 0

2 0

6 0

Z 2 0

ilu

a.

OL

20

20

• n

. 178

I

. 21

n

1

7

usi W fi

35

| 3 J 0.

SNIFFING

HEAD-DARTING

m

4 1 1 3

0 40

* 22 n 13 •

6 [~I • .

 1 n I

ji. • 1 .1 HI

9

cf

1

n

17

58

1

1

n

i

•

0 :

18

14

 4 -

15

head forefoot breast chest midback crest tailbase tail hmdfoot

.3. A J .

HEAD-DARTING rf

37 ^

• j . l 3 A.• I

SNIFFING 9

15

ii

mouth eye cheek ear ear ear ear mask head

nose • forehead • in front • behind belc

B
74 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

parts (Figure 37B,C). Based on a limited number of

observations, the nose, cheek, and ear received the

majority of nasal contacts to the head. Sniffing targets of the hindquarters differed only slightly between the two subjects. Targets in this area on the

female can be ranked in decreasing frequency from

scent-gland, under-tail, to anus. On the male's body,

the order of the first two targets was the same, the

scrotum was the third, and the anus was sniffed

least commonly.

As in Genetta the companion's head was the

most common target of head-darting, but in addition the neck, foreleg, and hindleg were also hit.

While these areas were also struck in the genets, the

relative proportion of darts to them was less, particularly in the case of the hindleg (Figure 36).

There is an obvious difference in targets between

animals. The male darted primarily at the neck,

foreleg, and hindleg (totaling 80% of 21 darts), and

the head and side were infrequently struck. The

female directed over 65% of 262 darts to the male's

head and struck at the neck less than 15% of the

time. The neck, hindleg, and foreleg were the next

most common targets (Figure 37A). Thirty percent

of the male's darts directed to the head struck the

ear region, while the mask and cheek were less

common targets.

In comparison with Genetta several quantitative

differences can be cited. Whereas the genets sniffed

the tail more often than the anogenital region (71%

of all sniffs to both areas), the civets exhibited only

a slight bias in favor of the tail (54% and 63% in

the male and female). In Genetta 33% of all sniffs

to the tail were directed to the base while in the

civets 54% of all such sniffs were aimed at the base

(male = 48%, female = 65%). Thus in Civettictis

the proximal portion of the tail and the anogenital

region seem to be more preferred or accessible

targets for sniffing. From the standpoint of hair

patterning these areas are quite distinctive (Figure

37c). The scent gland is a visually and physically

prominent attribute comparable in its alternating

pattern of white and black to the sides of the neck.

The tail of Civettictis differs from Genetta in that

most of the white hairs are concentrated on the

sides and ventral surface of the base. It is also

significant that the proportionately shorter tail of

the African civet provides a relatively smaller

sniffing area than in the genets.

Interaction through Sniffing and Head-darting:

the Sniff-Dart Axis

As sniffing and head-darting are prominent and

causally linked elements of interaction, it is important to consider some aspects of their occurrence.

The following analysis is based on the assumption

that head-darting occurs predominantly as a response to a proximate animal's sniffing or attempting to sniff its companion's body.

ROLES OF "SNIFFERS" AND "DARTERS".—Table 12

reveals that if a series of encounters is considered

the animal with the highest sniffing score may or

may not have the highest darting score. The meaning of these summated values, though, is difficult

to interpret because the relationship between two

animals can change during a series of encounters.

A number of patterns emerged, however, when

head-darting frequencies were compared for all

single encounters in which a given animal was the

sniffer. Three of the six animals exhibited statistically significant differences (Appendix 4). Animal

K, for example, darted significantly less frequently

than his companions when he dominated sniffing

(P <0.005). Animal C also darted less during encounters with F (P <.O5). There was no significant

difference in her darting frequency, however, when

these encounters are combined with the other ones

in which she was the predominant sniffer. Animal

O, on the other hand, darted more frequently in

encounters even though he was the dominant

sniffer (P = 0.025). The remaining three animals

displayed no consistent tendency to dart more or

less than their companions. Thus, while there is

usually a disparity in any given encounter in the

relative frequencies of sniffing and darting, some

sniffers predictably darted either more (O) or less

(K, C) than their companions. In I, F, and Mo

the level of darting bore no consistent relationship

to frequency of sniffing. Similar results were found

in the two African civets. The male, who dominated sniffing in eight of nine encounters, darted

significantly less than the female (P<0.01,

Appendix 5).

Appendix 6 records the darting scores of both

animals in 21 encounters in which only one genet

sniffed its companion. In 15 out of 21 cases the

sniffer received darts but did not dart itself. In the

remaining 6 encounters, both sniffer and nonsniffer

darted and the nonsniffer predominated. There ap-
NUMBER 239 75

pears to be a tendency for the sniffer to refrain from

darting, but this is not a statistically significant

pattern (P >0.05). On the other hand, of 68 encounters in which both animals sniffed, on only

10 occasions (14.7%) did the less sniffed animal

refrain from darting. When the less sniffed animal does dart, the level of darting varies individually both within and between encounters. These

findings tentatively suggest that the sniffer refrains

from darting as long as the companion confines its

activity to darting but does not smell back.

INDIVIDUAL VARIATION OF TARGET SELECTION.—

The overall distribution of sniffs and head-darts to

body targets may not be indicative of the specific

distribution exhibited by a given animal. Also,

the sniffing and head-darting orientation of any one

animal may or may not vary with different companions. To test the hypothesis that target distribution was independent of introduction of different companions, sniffing and head-darting targets

were grouped into four arbitrary categories: (1)

head and neck, (2) trunk (including shoulders,

chest, spinal crest, midback, rump, anogenital

region, and belly), (3) legs (fore- and hindlegs and

feet), and (4) tail (including tailbase); frequency

tabulations were ranked for these four body regions

on each companion that was encountered by a

given animal; and the results were tested with the

Friedman two-way analysis of variance (Appendices

7 and 8).

Though the head and neck were the predominant

targets, the absence of scores for the other regions

resulted in rank ties that produced insignificant

variance. The null hypothesis for sniffing was rejected at a probability of <0.017 for four of the six

animals (K, C, I, and O). For head-darting only

animals C and I deviated from a random variance

(P< 0.054). These results indicate that though most

animals darted and sniffed more often at certain

regions than others, the relative frequencies to these

regions were not necessarily consistent with different

companions. Thus, while almost all animals exhibited disproportionate contact distributions to their

companion's body regions only two of the six animals were inconsistent in the way they sniffed their

companions, and only two were consistent in headdarting at their companions.

SNIFF-DART RATIO AND TARGET SELECTION.—

Tables 17 and 18 summarize the relationship of

head-darting to sniffing and sniffing targets in

Genetta and Civettictis. The frequencies of these

two patterns were converted to rates to facilitate

comparison. Though the rates varied between encounters, the overall sniffing rates of the civets were

comparable to the higher rates shown by the

genets. The overall darting rate in the female civet

was two times greater than the highest rate displayed by a genet; the male displayed a darting rate

comparable to the lower rates in the genets. On

the average, then, the civets sniffed and darted at

relatively high rates. It is also evident that the

variation in sniff-dart ratios is not correlated with

any fluctuation in overall rates of sniffing and darting during an encounter or series of encounters.

It can be hypothesized that the distribution of

sniffs to the body parts is partially determined by

the relative proportion of sniffs that are responded

to with darts (i.e., the sniff-dart ratio). As headdarts usually cause a sniffing animal to desist (at

least momentarily), the predominance of tail and

hindbody sniffing appeared to be a strategy permitting maximum olfactory sampling at a minimum

risk of sustaining a dart.

To test this hypothesis, the body was divided into

three sections that maximally reflected the longitudinal polarity of sniffing distribution (Tables 17

and 18) and the frequency distribution of sniffs to

these three body parts was tested for low and high

sniff:dart ratio encounters with the Friedman twoway analysis of variance (Appendices 9 and 10). In

genets exhibiting sniff-dart ratios < 1:0.5, there was a

highly significant variance of the sniff distribution

to the three regions (P<0.001). When the same

genets had sniff:dart ratios > 1:0.33, the variance

between body regions did not differ from random.

Thus, though both groups showed an overall increasing gradient of sniffs from fore- to hindbody,

the pattern was consistent only in encounter series

characterized by low sniff:dart ratios.

These results are at odds with the prediction

that high levels of darting restricted sniffing to the

hindquarters. On the contrary, low sniff:dart

ratios do not predictably indicate a sniffing bias

to the hindbody, and genets strongly inclined to

sniff the fore- and midbody did so in spite of the

frequent darts they received. It appears that

although head-darting is often elicited by sniffing,

it does not predictably constrain the companion's

sniffing to the rear end.

In the African civets, sniffing variance between
76 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TABLE 17.—Relationship of sniffing to head-darting frequency and to sniffing

body targets (percentage in parentheses) in Genetta tigrina

1

2

3

4

5

6

7

8

9

10

11

Animal

Pair

F

0

C

K

Mo

K

I

K

C

I

C

Mo

C

F

I

0

C

0

o"- •

o" • •

9-.

rf..

d•'.'.

d'.\

2..

2..

2..

2..

2..

2 -

9 ••

I ?..

Mo 2 ••

I

F

9 ••

Sniff

Total

401

444

233

161

23G

218

160

92

58

60

58

64

79

1

0

43

5

38

8

30

2

1

per

10 min.

10.4

11.5

6.6

4.6

7.7

7.1

5.0

2.9

2.6

2.6

1.4

1.5

2.4

0.3

1.2

0.2

1.1

0.2

0.7

<0.1

<0.1

Head-dart

Total

31

62

170

42

16

8

24

8

49

4

17

21

0

57

9

30

33

4

4

7

12

2

per

10 min.

0.8

1.6

4.8

1.2

0.5

0.3

0.7

0.3

2.2

0.2

0.4

0.5

1.7

0.3

0.9

0.9

0.1

0.9

0.2

0.3

Sniff:Dart

ratio!

1:0.15

1:0.07

1:0.18

1:1.10

1:0.03

1:0.07

1:0.05

1:0.26

1:0.07

1:0.82

1:0.36

1:0.26

1:0.72

1:0.21

1:0.80

1:0.86

1:0.87

1:0.13

1:2.00

1:6.00

Sniffing

Forebody

129

65

44

24

54

47

30

21

15

18

15

25

6

0

0

3

2

11

6

5

1

0

(32.3)

(14.6)

(18.8)

(15.0)

(22.7)

(21.6)

(18.9)

(22.8)

(26.3)

(30.0)

(25.9)

(39.1)

(7.6)

(6^9)

(28.9)

(75.0)

(16.6)

--

Body Targets2

Midbody

149

137

58

50

52

44

60

33

12

19

9

6

9

1

0

3

3

2

2

5

0

0

(37.3)

(30.8)

(24.9)

(31.3)

(21.8)

(20.3)

(65.2)

(35.8)

(21.1)

(31.6)

(15.5)

(9.4)

(11.4)

(6^9)

(5~3)

(15.0)

(16.6)

--

Hindbody

122

242

131

86

132

126

68

38

30

23

34

33

64

0

0

37

0

25

0

20

1

1

(30.5)

(54.5)

(56.2)

(53.7)

(55.5)

(58.1)

(73.9)

(41.3)

(52.6)

(38.3)

(58.6)

(51.6)

(81.8)

(86.0)

(65.8)

(66.6)

--

Total duration

of encounters

per pair

(hr :min:sec)

6:24:06

5:52:57

5:08:39

5:18:24

3:57:25

6:41:20

5:29:40

5:54:02

5:34:16

6:45:39

5:52:44

Ratio of the number of companion-directed sniffs to the number of head-darts received from the companion.

2

 Forebody = head and neck; midbody = forelimbs, chest, shoulder, side, back, spinal crest, and belly; hindbody

tail , rump, hindlimbs, anogenital region. Percentages calculated only when sniffing frequency ^30.

the three body regions was statistically significant

in situations of high and low sniff:dart ratios

(P<.05), but a distinct difference in rank order

was exhibited. Though the hindbody was usually

the most frequently sniffed region in both groups

(Appendix 10), the forebody ranked second highest

in high ratio encounters and lowest in low ratio

encounters. So while there was no difference in

variance between the two conditions, the forebody

was a high frequency target more often when

sniff-dart ratios were high. In view of the limitations of the small number of animals, the data do

not lend themselves to further interpretation.

DISCUSSION

Vocalizations among Viverrids

Table 11 reveals that there is relatively little

variation between species in the number of calls and

their distribution within basic call groups. The

most widespread calls, namely hissing, whining,

screaming, and growling are known to occur in

several other mammalian orders. Whining, a characteristic call of infants given under a broad variety

of circumstances, is probably homologous to the

tremelo groan of Fossa and the bleating call in
NUMBER 239 77

TABLE 18.—Relationship of sniffing to head-darting frequency and to sniffing

body targets (percentage in parentheses) in Civettictis civetta

Encounter

and animal

1

2

3

4

5

6

7

8

9

<f

2

<f

2

<f

2

<t

2

<S

2

<f

2

d

2

<S

2

<t

2

TOTAL J

 ?

Total

105

69

46

20

22

3

8

4

23

23

15

3

45

23

16

1

2

18

282

164

Sniff

per

10 min.

23.8

15.6

13.2

5.7

4.0

0.5

1.2

0.7

6.6

6.6

4.3

0.9

11.1

5.8

3.2

0.1

0.5

4.2

7.2

4.2

Head-dart

Total

4

37

10

38

1

38

3

36

2

55

2

27

4

25

0

109

0

16

26

381

per

10 min.

0.9

8.4

2.4

10.9

0.2

7.0

0.4

6.1

0.5

15.8

0.6

8.1

0.5

6.5

22.5

3.8

0.7

9.9

Sniff:Dart

ratio*

1:0.35

1:0.05

1:0.82

1:0.50

1:1.72

1:0.33

1:4.50

1:0.75

1:2.39

1:0.08

1:1.80

1:0.66

1:0.55

1:0.17

1:6.81

1:8.00

1:1.35

1:0.15

Forebody

18

19

1

0

0

0

0

0

6

11

2

2

2

3

0

0

1

2

29

37

(26.3)

(32.7)

--

—

—

--

—

(4.8)

—

—

(13.8)

(29.1)

Sniffing Targets2

Midbody

18

15

10

3

7

0

2

0

5

1

1

0

11

1

1

0

0

1

55

21

(26.8)

(25.8)

—

—

__

—

—

(26.8)

—

—

(26.3)

(16.5)

Hindbody

31

24

18

12

10

1

4

3

11

4

9

1

28

13

14

1

0

10

125

69

(46.3)

(41.4)

—

--

--

--

—

(68.3)

—

—

(59.3)

(54.3)

Duration

(Min:sec)

44:00

34:39

53:54

59:24

36:30

33:15

38:30

48:24

40:26

388:22

* Ratio of the number of companion-directed sniffs to the number of head-darts received from the companion.

2

 Forebody = head and neck; midbody = forelimbs, chest, shoulder, side, back, spinal crest, and belly;

hindbody = tail , rump, hindlimbs (anogenital region excluded). Percentages calculated only when

sniffing frequency >_ 30.

Entries in columns 2 and 4 are means.

Nandinia. All are emitted repeatedly as responses

to temperature changes, hunger, and other sources

of discomfort or pain. Nandinia differs most from

the other species in retaining the call into adulthood, when it is produced in response to tactile

stimulation and apparently mild pain.

Low intensity whines are occasionally produced

by adult genets and African civets, but screaming is

the call that is usually emitted. Unlike whining,

screaming is a response to specific painful stimuli

that often arise during interaction with other

animals. It appears that during development

whining becomes decreasingly responsive to general

discomfort stimuli and is almost always exhibited

in an intense form (as screaming) to painful stimuli.

From the standpoint of avoidance context, hissing

and snorting are essentially identical calls. In all

species they are often associated with head-darting,

but in Paguma this connection appears to be obligatory. Humming seems to be a specialized derivative of growling. While long duration is shared by

both calls, humming is frequently modulated and

the species that display it do not growl. Though

humming was usually not produced under circumstances conducive to growling, when disturbed from

rest the Paguma often hummed after a series of
78

snorts. In Nandinia, animals mimiced the gradual

frequency oscillations of one another, but the function of the call is obscure.

Hooting and neighing are unique to the two

palm civets (Nandinia and Paguma) and no repetitive sound having equivalent intensity has been

heard in any other species. Eisenberg (pers. comm.)

heard a whinnying call in wild Ceylonese Paradoxurus that approximates my description of

neighing in Paguma. All three of these species are

solitary arboreal omnivores with a strong predisposition for fruit. It is feasible that these calls

function in spacing, for their brief, loud, and

repetitive sound properties make them easily localized over relatively long distances (Whitfield, 1971).

Though these three species of palm civets also

have perineal scent glands, an examination of

animals and museum specimens indicated low secretory rates of relatively weak scents. If it is

assumed that the paradoxurines evolved from semiarboreal ancestors in which scent markings served

a spacing function, it is conceivable that high

intensity calls have assumed at least some presumptive scent functions. The advantage of sound communication in a "volume environment" is that it

can be broadcast over long distances, its transmission is dependent on and enhanced by air current,

and it is doubtful that all parts of civets' living

volume are equally accessible for scent sampling.

It is curious that none of the terrestrial foraging

civets (Genetta, Civettictis, Fossa) have repetitive

high intensity calls in their repertoires. These calls

occur predominantly in small arboreal mammals

(galagos, tree hyraxes, monkeys), and terrestrial

species that are large (felids), or medium-sized and

social (canids). This pattern of occurrence suggests

that such a call might make a small, ground-living

species more vulnerable to predation.

In terrestrial species scent would seem to be a

more effective means of distance communication

because (1) the probability of a strange animal

entering the active space of a scent mark would be

greater in a two-dimensional than three dimensional

environment, and (2) scent marks do not betray

an animal's immediate whereabouts.

Coughing vocalizations were exhibited only by

the terrestrial foraging species. While the physical

properties of this call are probably quite similar

between species, the temporal patterning is distinctive. Coughs in Genetta and Fossa are emitted

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

singly or in volleys, and the interval distribution is

dispersed over a wide range of time (<l->6 0 sec).

In Civettictis and probably Viverricula, individual

coughs are almost always temporally clustered into

bouts composed of 5-6 elements, and these volleys

are usually separated from others by relatively long

periods of time (>60 sec).

In Civettictis, Viverricula, and Genetta, coughing

is probably homologous. In all three species it

clearly results from respiratory hesitation often

associated with sniffing, and stance and locomotory

transitions. In the adult African civets it was infrequently exchanged between animals. In motherinfant associations, however, it clearly functions as

a contact call: "the call is most frequently heard

from kittens and is given by an individual who

finds himself alone. The litter-mate at once responds

by repeating the call and moving to the caller . . .

the response of young kittens is completely automatic and invariable, and I never heard a call that

failed to evoke a response" (R. F. Ewer, pers.

comm.). The contingency analysis of this study

revealed that the callers were approached or sniffed

12%-20% of the times they coughed. Although

responsiveness to the call does not disappear, Ewer

noted that it gradually becomes more and more

conditional upon other factors in the situation

(Ewer and Wemmer, 1974).

The presumptive stimuli evoking this call are

always present, but animals do not cough continuously. Certain conditions appear to lower the threshold for calling. Confinement in an unfamiliar area

and separation from other animals or the keeper are

common conditions that elicit this call. When an

animal is sufficiently aroused, the coughing may

be spontaneous and bear no clear relationship to

ongoing activities. Prior to copulation, for example,

male genets coughed repeatedly while following

the female.

Coughing in volleys probably arises from a tendency for the glottis to close immediately after a

single cough and then snap open again. In Genetta

coughs are usually linked with expiration one for

one, but doublets and triplets also occur during

single expirations. The expulsion of volleys of

coughs during expiration in Civettictis could have

arisen from the doublet and triplet condition seen

in Genetta. Temporally dispersed coughing then

probably represents the primitive condition.

Coughing in Fossa resembles that of Genetta,
NUMBER 239 79

except that it can be evoked by human imitation

(Wemmer, 1971). It is probable that the call arose

independently in this species. Curiously, like Civettictis, Fossa is strictly a ground-living species.

Nonvocal Interaction in Genetta and Civettictis

A large number of behavior patterns are shared

by both Genetta and Civettictis; a small number are

different. The differences are best elucidated by a

consideration of timing and spacing. In Genetta

back-arching, crouching, and tail-flaring are associated with mutual or unidirectional visual fixation;

they occur in the companion's proximity, but out

of his reach. Back-arching and crouching are characterized by the tonic retention of a lateral or

frontal orientation to the other animal. During

minimal contact, sniffing may be unidirectional or

reciprocal; sniffing and head-darting are frequently

distinct roles, and mutual head-darting is rare.

Interaction is terminated by the departure of one

or both animals.

In Civettictis, back-arching and tail-flaring are

absent, and crouching is infrequent. Visual fixation

thus is not specified by tonic postural variants.

Minimal contact consists of mutual sniffing, and

sniffing and head-darting; unlike Genetta, reciprocal head-darting (sparring) is also common. The

recipient of darting frequently assumes (1) a lateral

or tangential orientation to the companion, with

the face deflected away, and/or (2) a reclining position on the ground. After the delivery of additional

head-darts, the episode is terminated by the darting

animal's departure.

Lateral or tangential body orientation to the

companion is a common component of contact

interaction in canids and has been referred to as the

T-position (Golani and Mendelssohn, 1971) or

scruff-shoulder orientation (Fox, 1971). In the

golden jackal, for example, the lateral animal is

the contact recipient; the companion may sniff, paw

at, or bite him, or may rest his head or forelegs on

the lateral animal's back. The T-position terminates

with circling or genital licking (Golani and

Mendelssohn, 1971). In the wolf, dog, and coyote,

a reclining position may be assumed if the lateral

animal is pushed (Fox, 1971). Lateral recumbency

(passive submission: Schenkel, 1967) is also a reaction to being sniffed in the genital region. According to Schenkel (1967:324) the most obvious

characteristic of the recumbent animal is "the readiness to actively enter into contact with the superior"

animal; the reclining animal may whine, make

licking movements, push with its nose, and wag the

tail. In reclining civets, only the head and neck

are moved in response to the proximity and headdarts of the standing animal; motion only appears

to provoke further darting. Nevertheless, while

types of contact differ, Civettictis is basically similar

to the canids in that contact is received during

lateral orientation and recumbency is a response to

contact.

While there are individual differences in the

orientation of sniffs and darts, in both Genetta and

Civettictis there is a general trend for the forebody

to be the darting target, and the fore- and hindbody

to be sniffing targets. Though other workers have

not quantitively documented observations of oriented contact, it has been suggested that certain

body markings serve as contact targets (Fox, 1969;

Kleiman, 1967).

The body targets of sniffing and darting are the

same in these two species, but the hair patterning

differs. There is thus an indication that white body

markings may serve several different functions. Presumptive general functions can be deduced by

evaluating (1) whether the marking is always prominent or can be hidden, (2) whether the size, proportions or shape of the design is strongly influenced

by body configuration, and (3) whether the marking

is characteristically oriented to companions or

oriented to by companions. The contrasting markings on the bodies of Genetta and Civettictis do not

exhibit these characters equally. In the African

civet, the white spinal stripe is visible only during

crest erection and the white perineal gland is often

concealed by the tail; only the gland receives

oriented contact. Neither marking's configuration is

altered significantly when viewed from different

positions.

The design properties of the eye spots and tail of

Genetta and the face and neck markings of Civettictis, however, alter considerably during positional

or postural changes in relation to the viewer. Only

in Civettictis, though, is there a correspondence

between the location of contrast patterns and

oriented contact. On the basis of these different

properties, contrast markings may (1) enhance

visibility of the body or its movements in animals

that are nearby but out of contact range, (2) specify
80 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

postural configurations of the body or the part of

the body carrying the mark, and (3) specify targets

for companion-oriented contact.

The following outline summarizes some of the

possible changes associated with the transition

from a genet-like grade of behavioral organization

to that exhibited by the African civet. It is based

on the assumption that Genetta is a relatively

primitive viverrine and that Civettictis evolved

from an ancestor resembling Genetta. Items are

numbered for convenience, but the separation does

not imply a lack of causal relationship between

them.

1. Relative shortening of tail length and degeneration of its highly patterned design; loss of tailflaring capacity.

2. Loss of ability to maintain protracted archedback stances; compensatory development of the

spinal crest but retention of the tendency to

maintain a lateral body orientation to other animals during maximal crest erection.

3. Origin of lateral body orientation with neck

deflection, and its incorporation with reclining

body positions as intermediary components in the

sniff-dart axis; concomitant elaboration of highly

patterned head and neck markings from a basic

genet-like design.

4. Enlargement of the perineal gland and elaboration of its coloration; derivation of the normal

quadrupedal stance from a squatting or reversed

quadrupedal upright; specialization of Flehmen.

The genet and African civet are more similar

behaviorally to one another than to other species

of carnivores. While lateral orientation and reclining are also components of contact interaction in

canids, it is probably a simplification to view

Civettictis as a more social species. There is no

evidence suggesting it is anything but solitary. Perhaps the exigencies of a totally terrestial existence

select for an interaction design that permits the

establishing of social roles in minimal time. It is

conceivable that the assumption of oriented stances

in the African civet facilitates transition from

sniffing and darting to maximal contact interaction.

Addendum

Shortly after this manuscript was revised for publication two important papers

appeared dealing with the natural history and behavior of the small-spotted

genet, Genetta genetta, in Europe. They are B. Gangloff and P. Ropartz,

Le Repertoire Comportemental de la Genetta, Genetta genetta (Linn6), La

Terre et la Vie (1972) 26(4):489-560; and M. Delibes, Sobre Alimentacion y

Biologia de la Geneta (Genetta genetta L.) en Espana, Donana, Acta Vertebrata

(1974) 1:143-199.

Gangloff and Ropartz' study is a lengthy description of the social, nonsocial,

and predatory behavior of captive animals. It is evident that the social behavior

of this species is qualitatively similar or identical to that of Genetta tigrina;

however, quantitative data on social interaction are not presented on which to

base further comparisons. Delibes' analysis of feeding habits reveals the preponderance of small vertebrates and insects in this species' diet. This is the only

large contribution available on this aspect of genet ecology.
Appendices

ENCOUNTER SCHEDULES AND STATISTICAL PROCEDURES

Appendix 1:

Encounter

number Date

1

2

3

4

5

6

7

8

9

10

1

2

3

4

5

6

7

1

2

3

4

5

6

7

8

9

10

1

2

3

4

6Sep69

8 Sep69

10 Sep 69

12 Sep 69

14 Sep 69

16 Sep 69

18 Sep 69

20 Sep 69

22 Sep 69

24 Sep 69

Encounter Schedules for Genetta

First

animal

introduced

Duration in

cage prior to

encounter

(hrs:min)

MALE-MALE ENCOUNTERS: F AND O

F

O

F

O

F

O

F

O

F

O

3:30

3:50

3:50

4:10

3:45

3:25

4:00

4:15

4:10

4:20

Second

animal

introduced

1900

1940

1915

1910

1645

1630

2010

1935

1945

1930

FEMALE-FEMALE ENCOUNTERS: C AND I (familiar)

20 Jun 69

9 Jul 69

17 Jul 69

25 Jul 69

31 Jul 69

7 Aug 69

15 Aug 69

I

I

I

C

c

I

I

4:15

—

3:50

3:45

2:45

3:40

2:05

2030

1900

2010

1930

1900

2040

2000

FEMALE-FEMALE ENCOUNTERS: C AND MO

18 Aug 69

20 Aug 69

22 Aug 69

24 Aug 69

26 Aug 69

28 Aug 69

30 Aug 69

1 Sep 69

3 Sep 69

5 Sep 69

25 Sep 69

27 Sep 69

28 Sep 69

1 Oct 69

c

Mo

C

Mo

C

Mo

C

Mo

C

Mo

4:30

3:15

1:15

2:30

4:47

2:45

3:12

4:05

3:50

4:50

2130

1945

1630

1845

1855

1930

1930

2000

1910

2020

FEMALE-FEMALE ENCOUNTERS: I AND MO

Mo

I

Mo

I

3:20

4:25

3:00

3:00

1950

2050

1905

1900

tigrina

Encounter

duration

(min:sec)

52:53

45:47

31:24

34:20

34:09

33:35

40:24

46:10

31:39

33:45

30:58

33:44

32:56

32:22

32:55

33:11

31:19

34:57

40:50

36:31

33:34

34:45

49:22

46:15

35:55

39:01

50:10

35:25

37:17

37:27

41:01

81
82

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Appendix 1—Continued

Encounter

number

5

6

7

8

9

10

Date

3 Oct 69

5 Oct 69

7 Oct 69

9 Oct 69

11 Oct 69

IS Oct 69

First

animal

introduced

Mo

1

Mo

1

Mo

I

Duration m

cage prior to

encounter

(hrs:min)

3:35

2:40

3:00

3:20

3:30

3:30

Second

animal

introduced

1915

1930

1615

1940

1630

1930

Encounter

duration

(min:sec)

45:19

39:42

45:23

48:38

36:34

38:53

MALE-FEMALE ENCOUNTERS: K AND MO

1

2

3

4

5

6

7

8

9

10

1

2

3

4

5

6

7

8

9

10

1

2

3

4

5

6

7

8

9

10

1

2

3

4

5

6

9 Jun 69

16 Jun 69

23 Jun 69

30 Jun 69

7 Jul 69

14 Jul69

21 Jul 69

28 Jul 69

4 Aug 69

11 Aug 69

Mo

Mo

K

Mo

K

Mo

Mo

K

Mo

K

4:00

3:10

3:30

3:50

3:45

3:00

2:25

6:50

2:15

3:15

2100

2010

2030

1620

1915

1930

1715

1630

1915

1930

MALE-FEMALE ENCOUNTERS: K AND C (familiar)

14 Oct 69

16 Oct 69

18 Oct 69

20 Oct 69

22 Oct 69

24 Oct 69

26 Oct 69

28 Oct 69

30 Oct 69

1 Nov 69

K

C

K

C

K

C

K

C

K

C

3:00

3:40

3:40

4:05

3:05

3:25

3:30

4:00

3:20

4:05

MALE-FEMALE ENCOUNTERS: F AND I

2 Nov 69

4 Nov 69

6 Nov 69

8 Nov 69

10 Nov 69

12 Nov 69

14 Nov 69

16 Nov 69

18 Nov 69

20 Nov 69

F

I

F

I

F

I

F

I

F

I

3:45

2:50

6:20

3:00

3:15

3:22

3:50

3:25

5:05

4:30

MALE-FEMALE ENCOUNTERS: O AND C

21 Nov 69

23 Nov 69

25 Nov 69

27 Nov 69

29 Nov 69

1 Dec 69

O

C

o

c

o

c

3:00

2:55

2:55

4:10

3:48

3:00

1635

1940

2030

2050

1940

2040

2000

2000

1940

2010

1925

1620

1950

1615

1645

1652

1940

1655

2030

2100

1850

1625

1840

1740

1658

1920

29:54

30:06

31:22

36:16

32:11

33:06

22:32

27:04

32:49

33:19

35:14

35:44

34:04

36:21

36:11

35:31

39:47

34:24

33:44

31:57

39:10

33:32

36:36

31:16

33:06

39:48

34:14

33:14

38:38

33:10

39:32

36:39

34:55

31:39

30:23

33:56
NUMBER 239

7

8

9

10

3 Dec 69

5 Dec 69

7 Dec 69

9 Dec 69

O

C

O

C

3:55

4:00

3:00

4:20

2015

1930

1630

2020

28:11

32:18

35:24

31:19

83

MALE-FEMALE ENCOUNTER: F AND M O

11 Dec 69 F 3:00

MALE-FEMALE ENCOUNTERS: O AND I

1600 11:40

1

2

3

4

5

6

7

8

9

10

1

2

3

4

5

6

7

8

9

10

1

2

3

4

5

6

7

8

9

10

30 Dec 69

1 Jan 70

3 Jan 70

5 Jan 70

7 Jan 70

9 Jan 70

11 Jan 70

13 Jan 70

15 Jan 70

17 Jan 70

o

i

o

I

o

I

o

I

o

I

3:30

3:20

3:35

4:05

3:50

4:50

2:40

3:40

3:00

3:10

1930

1905

1935

2040

1950

2050

1940

1940

1900

1610

MALE-FEMALE ENCOUNTERS: F AND C

18 Jan 70

20 Jan 70

22 Jan 70

24 Jan 70

26 Jan 70

28 Jan 70

30 Jan 70

1 Feb 70

3 Feb 70

5 Feb 70

F

C

F

C

F

C

F

C

F

C

3:30

2:20

3:05

3:20

3:15

4:10

3:00

4:25

4:15

4:05

MALE-FEMALE ENCOUNTERS: K AND I

6 Feb 70

8 Feb 70

10 Feb 70

12 Feb 70

16 Feb 70

18 Feb 70

20 Feb 70

22 Feb 70

26 Feb 70

2 Mar 70

K

I

K

I

K

I

K

I

K

I

Appendix 2: Encounter

3:00

3:30

4:15

3:20

4:00

3:30

4:00

3:45

3:30

3:00

1915

1845

1930

1650

1845

2010

1845

2020

2015

2000

(familiar)

1945

2000

2015

1640

2000

2000

2000

1845

1630

1900

Schedules for Civettictis

32:14

34:44

52:50

32:54

35:08

33:16

33:01

34:16

32:43

32:56

32:11

32:58

35:40

34:42

29:45

32:27

28:20

33:35

36:03

33:59

32:57

27:21

30:32

29:42

35:20

33:30

33:27

36:27

29:56

29:12

civetta

Encounter

number

1

2

3

4

(male-female encounters)

Date

2 Nov 70

3 Nov 70

4 Nov 70

5 Nov 70

Encounter

onset time

1500

1530

1445

1500

Encounter

duration

(min:sec)

44:00

34:39

53:54

59:24
84 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Appendix 2—Continued

Encounter

number

5

6

7

8

9

Appendix 3:

Date

9 Nov 70

10 Nov 70

12 Nov 70

13 Nov 70

16 Nov 70

Statistical Procedures

Encounter

onset time

1530

1515

1500

1500

1530

for Tail-flaring

Encounter

duration

(min:sec)

36:30

33:15

38:30

48:24

40:25

Frequency

in Genetta tigrina

Wilcoxon matched-pairs, signed-ranks test; N = number of series encounters in which at least

one animal jaw-gaped and both animals tail-flared (see Table 13); variable: tail-flaring frequency; significance level (two-tailed): 0.05

Jaw-gaper

K

Mo

I

C

I

F

I

I

7

5

20

22

2

1

3

2

c

K

Mo

Mo

O

C

F

K

Companion

13

19

27

23

11

8

1

1

Rank Statistics

- 6

- 1 4

- 7

- 1

- 9

- 7

+2

+ 1

-4.0

-8.0

-5.5

-1.5

-7.0

-5.5

+3.0

+ 13

N

T

P

= 8

= 43

= >0.05

Appendix 4: Statistical Procedures for Head-darting Frequency

in Genetta tigrina (between Sniffers)

Wilcoxon matched-pairs, signed-ranks test; N = number of single encounters in which (1) each

animal sniffed its companion, and (2) at least one animal head-darted; sniffer = animal that

sniffed more frequently than its companion; variable: head-dart frequency; significance level: 0.05

Sniffer Sniffed Rank Statistics

K 0

8

0

0

7

3

0

0

2

C 33

15

20

13

3

9

0

0

8

I

Mo

K

I

Mo

3

37

9

4

32

6

4

1

5

9

5

5

4

0

1

8

2

4

- 3

- 2 9

- 9

- 4

-25

- 3

- 4

- 1

- 3

+24

+ 10

+ 15

+9

+3

+8

- 8

- 2

+ 4

- 3

- 9

- 7

-53

- 8

- 3

-53

- 1

- 3

+ 18

+ 14

+ 17

+ 12.5

+5

+ 10.5

-10.5

- 4

+6

N

T

P

= 9

= O

= <0.005
NUMBER 239 85

Sniffer Sniffed Rank Statistics

c

I

Mo

O

F

1

5

0

0

0

0

0

0

0

0

0

0

0

0

0

0

18

0

2

0

1

1

0

3

0

3

0

0

7

0

0

0

0

2

0

1

0

5

10

6

6

2

9

5

0

6

0

0

0

0

0

1

0

1

0

5

2

O

F

F

K

F

Mo

C

I

C

K

F

I

C

O

0

4

7

6

9

11

1

14

5

7

6

9

11

1

14

5

0

2

0

4

2

0

3

0

37

0

1

1

0

1

2

1

4

1

1

3

1

6

8

8

2

0

2

1

1

1

2

9

2

4

4

0

1

17

1

14

8

+ 1

+ 1

- 7

- 6

- 9

- 1 1

- 1

- 1 4

- 5

- 7

- 6

- 9

- 1 1

- 1

- 1 4

- 5

+ 18

- 2

+2

- 4

- 1

+ 1

- 3

+3

- 3 7

+3

- 1

- 1

+ 7

- 1

- 2

- 1

- 4

+ 1

- 1

- 2

- 1

- 1

+2

+2

+4

+2

+7

+4

- 1

+5

- 2

- 9

- 2

- 4

- 4

+ 1

- 1

- 1 6

- 1

- 9

- 6

+2

+2

- 9

- 8

-12 3

- 1 5

- 2

- 1 6

- 7

- 4

- 3

- 5

- 6

- 1

- 7

- 2

+8.0

-3.5

+3.5

-7.0

- 1 3

+ 13

- 5 3

+53

+9.0

+9.0

- 3 3

-3.5

+ 11.0

- 3 3

- 7 3

- 3 3

-10.0

+ 3 3

-3.5

- 7 3

-2.0

-2.0

+6.0

+6.0

+ 103

+6.0

+ 14.0

+ 103

-2.0

+ 13.0

-6.0

+ 15.0

-6.0

+ 103

+ 103

+2

- 2

- 6

- 2

- 5

- 4

N

T

P

N

T

P

N

T

P

N

T

P

N

T

P

N

T

P

= 18

= 54

= >0.05 (nj.)

- 7

= O

= 0.05

= 9

= 173

= >0.05 (n.$.)

= 11

= 233

= >0.05 (nj.)

= 15

= -24 *

= 0.025

= 6

= 2

= >0.05 (n.s.)

• The negative sign of the T value indicates statistical significance for the opposite hypothesis.
86 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Appendix 5: Statistical Procedures for Head-darting Frequency

in Civettictis civetta

Wilcoxon matched-pairs, signed-ranks test; N = number of encounters;

variable: head-dart frequency; significance level: 0.025

Sniffer Sniffed Rank Statistics

4

10

1

3

2

4

0

16

37

38

38

36

27

25

109

0

- 3 3

- 2 8

- 3 7

- 3 3

- 2 5

- 2 1

-109

+ 16

-5.5

-4.0

-7.0

-5.5

-3.0

-2.0

-8.0

+ 1.0

N

T

P

= 8

= 1

- <0.01 >0.005

Appendix 6: Statistical Procedures for Head-darting Frequency

in Genetta tigrina (between Sniffers and Nonsniffers)

Wilcoxon matched-pairs, signed-ranks test; N = number of single encounters in which only one

animal sniffed its companion, and at least one head-darted; variable: head-dart frequency; significance level: 0.05

Encounter

number

1

8

10

4

8

1

5

6

7

8

1

2

3

6

10

3

4

5

6

8

10

Nonsniffer

I

I

C

C

o

c

F

I

1

1

1

2

1

2

9

2

4

4

7

6

9

11

5

2

0

2

1

1

1

Sniffer

F

Mo

Mo

O

C

O

C

O

0

0

0

0

0

0

0

0

5

0

0

0

0

0

0

6

2

9

5

0

6

d

+ 1

+ 1

+ 1

+2

+ 1

+2

+9

+2

- 1

+4

+7

+6

+ 9

+ 11

+ 5

- 4

- 2

- 7

- 4

+ 1

- 5

R'

+ 2.0

+2.0

+2.0

+7.0

+2.0

+7.0

+ 13.5

+7.0

-2.0

+9.0

+ 12.0

+ 11.0

+ 13.5

+ 15.0

+ 10.0

- 5

- 2

- 6

- 3

+ 1

- 4

R"

+ 3.5

+3.5

+ 3.5

+8.5

+3.5

+8.5

+ 19.5

+8.5

-3.5

+ 12.0

+ 17.5

+ 16.0

+ 19.5

+21.0

+ 14.5

-12.0

-8.5

-17.5

-12.0

+3.5

-14.5

Statistics

N =

T --

P =

= 21

= 68.0

= >0.05

Appendix 7: Statistical Procedures for Testing Distribution of

Nasal Contacts to Four Body Regions in Genetta tigrina

Friedman two-way analysis of variance; conditions = 4 general body targets (I «• head and neck,

II = trunk, HI — legs, IV = tail); variable: nasal contact frequency; significance level: 0.05
NUMBER 239 87

Animals

encountered

Individual

C

I

Mo

Individual

K

Mo

F

O

I

Individual

Mo

F

C

K

Individual

I

C

K

F

Individual

F

I

C

Individual

I

O

C

Mo

K

C

I

Mo

O

F

N

24

21

47

44

14

6

2

15

6

0

18

30

5

25

54

1

65

3

11

0

129

0

1

/

Rank

2.0

2.0

2.0

~6lJ

2.0

3.0

2.0

3.0

3.0

Ts^o

4.0

2.0

3.0

2.0

IsTo

2.0

3.0

2.0

4.0

1L0

1.0

2.0

3.0

To

2.0

3.0

2.0

4.0

ffo"

N

57

40

79

52

12

18

3

24

2

0

29

59

7

10

74

0

125

5

1

0

151

1

0

Conditions

II

Rank

3.0

4.0

3.0

IThO

3.0

2.0

3.0

4.0

4.0

I61T

3.0

2.0

4.0

4.0

TsTi

3.0

2.0

3.0

2.0

loo

3.0

3.0

1.0

To

2.0

4.0

4.0

2.0

12J0

N

9

4

9

27

4

3

0

6

0

0

5

24

1

2

17

0

69

2

3

0

48

0

0

III

Rank

1.0

1.0

1.0

3^0"

1.0

1.0

1.0

1.5

1.0

5T

1.5

2.0

1.0

1.0

5T

1.0

1.0

1.0

2.0

5iO~

2.0

1.0

2.0

ITo"

2.0

2.0

2.0

2.0

8lf

N

70

26

82

97

28

52

0

12

0

1

8

45

17

27

93

0

185

33

23

1

72

0

0

IV

Rank

4.0

3.0

4.0

[To

4.0

4.0

4.0

1.5

2.0

T5J0

1.5

4.0

2.0

3.0

ioJ

4.0

4.0

4.0

2.0

liJO

4.0

4.0

4.0

12i0

4.0

1.0

2.0

2.0

~9J0

Statistics

Xr2 = 8.2

P = 0.017

Xr* = 6.6

P = <0.01

Xr1 = 11.7

P = <0.001

XrI = 6.3

P = 0.094

Xr» = 8.4

P = <0.017

Xr»= 1.5

P = 0.754

Appendix 8: Statistical Procedures for Testing the Distribution

of Head-darting to Four Body Regions in Genetta tigrina

Friedman two-way analysis of variance; conditions = 4 general body targets (I = head and

neck, II = trunk, III = legs, IV = tail); variable: head-darting frequency; significance

level: 0.05

Animals

encountered

Individual K

C

I

Mo

N

34

2

6

/

Rank

4.0

4.0

4.0

Tzo

N

1

0

0

Conditions

II

Rank

1.0

2.0

1.0

To

N

2

0

1

III

Rank

2.5

2.0

2.5

To

N

2

0

1

IV

Rank

2.5

2.0

2.5

To

Statistics

Xr* = 6.6

P = 0.075
88 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Appendix 8—Continued

Conditions

Animals

encountered

Individual C

K

Mo

F

I

Individual I

Mo

O

C

F

Individual Mo

I

C

K

Individual O

F

C

I

Individual F

I

O

C

N

135

16

28

4

3

12

20

7

7

20

6

31

4

12

2

23

36

Rank

4.0

4.0

4.0

4.0

1T6

4.0

4.0

4.0

4.0

I<ro

4.0

4.0

4.0

T2l)

4.0

4.0

4.0

Tzo

4.0

4.0

4.0

TiTo

N

12

0

0

2

0

0

0

0

0

0

0

5

0

0

0

1

0

Rank

3.0

2.0

2.0

3.0

1T0

2.0

2.0

2.0

2.0

To

2.0

1.5

1.0

TB '

2.0

2.0

2.0

To "

2.0

2.0

2.0

To ~

N

9

0

0

0

0

0

0

0

0

0

2

8

0

0

0

1

0

III

Rank

2.0

2.0

2.0

1.0

To

2.0

2.0

2.0

2.0

To"

2.0

1.5

2.5

To

3.0

2.0

2.0

To

2.0

2.0

2.0

To"

N

3

0

0

1

0

0

0

0

0

1

2

1

0

0

0

1

0

IV

Rank

1.0

2.0

2.0

2.0

To"

2.0

2.0

2.0

2.0

To"

2.0

3.0

2.5

TF

1.0

2.0

2.0

To

2.0

2.0

2.0

To"

Statistics

Xr1 = 8.1

P = 0.033

Xr1 = 7.2

P = 0.054

Xr» = 6.1

P - <0.148

Xr1 = 4.2

P = 0.148

Xr* = 5.4

P = 0.175

Appendix 9: Statistical Procedures for Testing the Distribution

of Nasal Contacts to Three Body Regions in Genetta tigrina

Friedman two-way analysis of variance; conditions = 3 general body targets (I = forebody,

II = midbody, III = hindbody); first test: encounter series in which the sniff-dart ratio was

< 1:0.5, second test: > 1:0.36; variable: nasal contact frequency, significance level: 0.05

Animal

and

companion

K

I

C

K

Mo

Mo

C

Mo

C

O

O

F

(I)

(*)

(K)

(Mo)

(K)

(I)

(Mo)

(Q

(I)

(I)

(

F

)

(O)

N

21

30

44

47

54

5

15

25

15

3

65

129.

I

Rank

1.0

1.0

1.0

2.0

2.0

1.5

2.0

2.0

2.0

1.5

1.0

2.0

191F

Conditions

N

33

60

58

44

52

5

9

6

12

3

137

149

II

Rank

2.0

2.0

2.0

1.0

1.0

1J

1.0

1.0

1.0

1.5

2.0

3.0

21.0

N

38

68

131

126

132

20

34

33

30

37

242

122

III

Rank

3.0

3.0

3.0

3.0

3.0

3.0

3.0

3.0

3.0

3.0

3.0

3.0

34.0

Statistic

< 1:0.5

Xr1 = 19.1

P = <0.001
NUMBER 239

K.

I

I

O

C

C

F

I

(Q

(Mo)

(C)

(C)

(O)

(F)

W

(F)

24

6

18

11

2

6

0

1

1.0

3.0

1.0

2.0

2.0

1.0

1.5

2.5

14.0

50

2

19

2

3

9

0

0

2.0

2.0

2.0

1.0

3.0

2.0

1.5

1.0

TilT

86

0

23

25

0

64

1

1

3.0

1.0

3.0

3.0

1.0

3.0

3.0

2.5

19.5

> 1:0.36

Xr1 = 2.3

P = <0.3

89

Appendix 10: Statistical Procedures for Testing the Distribution

of Nasal Contacts to Three Body Regions in Civettictis civetta

Friedman two-way analysis of variance; conditions = 3 general body targets (I = forebody,

II = midbody, III = hindbody); first test: encounter series in which the sniff-dart ratio was

< 1:0.5, second test: > 1:0.33; variable: nasal contact frequency; significance level: 0.05

Encounter

and

animal

2

2

3

4

4

5

6

6

7

8

9

1

1

3

5

7

$

9

S

$

9

3

o

$

&

S

s

9

9

9

9

N

1

0

0

0

0

6

2

2

2

0

1

18

19

0

11

3

Rank

1.0

1.0

1.0

1.0

1.5

2.0

2.0

3.0

1.0

1.0

3.0

1.5

2.0

1.5

3.0

2.0

Too

N

10

3

7

2

0

5

1

0

11

1

0

18

15

0

1

1

Conditions

II

Rank

2.0

2.0

2.0

2.0

1.5

1.0

1.0

1.0

2.0

2.0

1.5

nnr

1.5

1.0

1.5

1.0

1.0

To

N

34

17

11

6

3

12

11

1

32

15

0

62

29

1

8

17

III

Rank

3.0

3.0

3.0

3.0

3.0

3.0

3.0

2.0

3.0

3.0

1.5

3"blT

3.0

3.0

3.0

2.0

3.0

TO"

Statistics

< 1:0.5

df = 2

Xr1 = 9.8

P = <0.01

> 1:0.33

df = 2

Xr1 = 6.4

P - <0.05
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1966. An Introduction to the Osteology of Mammalia. 382

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1952. Vergleichende Studien an unseren Marderartigenvom "Drohschrei" und anderen Stimmlauten. Pirsch/

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1956. Das Verhalten der Katzen. Handbuch der Zoologie,

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1953. Comparative Studies on the Behaviour of Anatinae.

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1965. The Comfort Movements of Anatidae. Behaviour,

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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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1969. Notes on Breeding the African Civet Viverra civetta

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