Vol. 7, pp. 108-130

Asiatic Herpetological Research

1997

Behavior of Varanus griseus during Encounters with Conspecif ics

ALEXEY YU.TSELLARIUS AND ELENA YU.TSELLARIUS
Program VARAN, Dept. Herpetology, Zoological Institute. 199034. St. Petersburg. Russia
Abstract. - In 1990-1993 in the western region of the deserts of Kyzylkum (Uzbekistan), constant observations ofof desert monitors in natural conditions were conducted. Described are manifestations of variousagroupemotional states of monitors, and common types of monitor interaction. Given are detailed descriptions of thecourse of contacts between the animals, illustrated by photographs. Fights were rarely noted and only between
unfamiliar lizards.

Itisntualization of the fight.

not from a
proposed that ritual combat arises from displays of dominance andvariedandnot
The behavior of monitors during contacts is highly
stereotypical. Dataof a complex, mammal-like social structure
Considered are probable mechanisms of intraspecific communication of monitor lizards.

from observations

Key words:

attestssocial behavior, social relations. Reptilia. Sauria, Varanus griseus

Introduction
In the article atrevealthethe population.

into the existence

hand we have made an attempt tocommunicational

functionbehavioral acts of desert monitor.

ofcommonWe have made also

an attempt to describe some types of interactions
between monitors and the role of these interactions insocial organization of population.

We will not touch

Behavior of animals during intraspecific contacts is
usually considered as a succession of behavioral acts,
but not as a social interaction directed at the mainte-

nance of long-term interrelations within a socium.

The analysis of behavior is reduced to statistical analysis of the sequence of the behavioral acts oftants.

contac-Generally, the task of the analysis is limited toupon behavior connected with courtship and mating

discovery of the most probable response of a lizard to
In analysis one seldom
the acts of another lizard.

in the present article.

allows for the circumstances, under which the interac-

In 1989.

when a program for the study of endan-

gered species of reptiles was developed by us, theemphasis was put upon their ecology.
When we stumbled onto the fact, that the behavior ofprincipal

the monitor lizard

ismuch more complex than was

imagined, we no longer had the opportunity to alter
the program of study.
Hence, the gathering of etiological material had to be conducted at the same time
Although we are aware of theas the primary tasks.

incompleteness of our data and the necessarily schematic and fragmentary nature of our description, we
do not consider its publication to be in vain. Thisissue has not only theoretical but practical significance since desert monitor populations arouse serious

apprehension

in

many

regionsofMiddleAsia(Darevsky,Orlov 1988. Khodzhaev 1989)
There exists a vast literature dedicated to the ele-mentary behavioral acts of lizards (see the survey of
Carpenter. Ferguson 1977). The behavior of somespecies of monitor lizards has also been described insome detail (Auffenberg 1981a; 1981b: 1983: 1988;
Carter; 1990; Davis etal.

1986; Deraniyagala 1958;

Gaulke 1989; Horn et al. 1994; Mertens: 1946).

But

in the most of herpetological papers the social motives

and purposes of behavioral acts are not examined.

tion took place.

The history of interrelations of con-tactants, as a rule, is not taken into

Actually,thesocialinteractionisaccount at

all.

considered as aclosed system with internal self-regulation, independent of the structure of the socium.
Viability of the population (the ability for bal-

anced reproduction in particular) is to a considerable
extent determined by its social structure.

The social

structure of many species is rather impressed by envi-

ronmental conditions and the reptiles are not an
exception (Panov, Zykova, 1985; Poly nova, Panyushkin 1982; Polynova, 1990; Stamps; 1977). A social

response of population to environmental changes isoften species-specific (Polynova, 1990) and is notnecessarily adaptive (Plyusnin, 1990). Peculiarities of
response of socium to the external influences are
determined for the most part by two closely interdependent, species-specific systems of animal activity:a) a production of the signals which carry information

about the animal and its circumstances; b) a perception, processing and analysis of these signals in conjunction with others external and internal irritants.
The principles of organization of these processes may

be termed as a "language" and a "mentality" of a species. This issue is closely linked to the problem of1997

Asiatic Herpetological Research

management of populations and demands

Vol. 7, p. 109

study.

monitor lizard was specially caught, usually in themorning hours. In the evening of that same day we

The Region of Studies

attempted to establish from tracks the site whereanother monitor would spend the night. At dusk we

carefulThe studies were conducted in the western area of thesand deserts of Kyzylkum. Uzbekistan.

The coordi-

nates of our permanent camp were 40° 40' N and 62°
08' E.

The landscape in this region is typical of the--released the monitor captured earlier into a nearby
burrow.
In the morning, as the monitors emerged

from their overnight shelters, encounter was inevitaIt was only
possible to perform these operations
from time to time so as not to disturb the lizards andble.

Kyzylkumsandy ridges, bushy undergrowths(mainly Haloxylon persiewn, and Calligonum sp.),

disrupt the normal course of their lives.

and sparse grass (Care.x physodes predominates) (Fig.

instances discussed in this article, data from tracking

In

some

In the region of studies there is a rather high abun-

was also used when it was possible to precisely anddance of rodents, for the most part Rhombomys opi-

fully reconstruct the course of events from the tracks.

mus

During observations from hiding, detailed stenographic notes were taken which were deciphered

1

).

and

Both

Spermophylopsis leptodactylus.
inhabited and abandoned colonies of Rh. opiums—

with their complex system of underground passages
and location not farther than 150-200 meters from one

another -- serve the monitors as refuges and hunting

immediately after observations were concluded.
Some of the encounters were photographed (from 5 to
16 frames per encounter). Unfortunately, a lack ofgrounds.

means prevented us from employing photographic
documentation to the extent that was necessary. In all.

Materials and Methods

we observed 37 instances of contact between males, 8
instances

Observations were conducted from 1990 to 1993 forperiods of four to seven months annually.

In

contact

betweenfemales,

and

21

all,

observations were made for a duration of more than

20 months.

ofinstances of contact between the sexes.

beginning of April, the period inwhich monitor lizards come out of hibernation, they

Results

In thewerecaught, measured, weighed, marked, andreleased at the site of their capture - usually in thecourse of a few hours after being caught.
specialA

mark permitted

us to identify the animals by theirtracks (Tsellarius and Cherlin, 1991 ).

Henceforth, forthe greater part of the season of monitor lizard activity

The Spatial Structure of the Population and
Annual Dynamics of Activity
Data pertaining to the spatial structure of the population is currently being readied for publication in a separate article.

Here, however, it is only necessary toaddress a few words to this issue.

The number of

we conducted continuous observations of the animals

monitor lizards in the region of studies consisted on— following their tracks and observing them visually

the average of four adult individuals per square kilo-

from camouflaged holes situated in the places where
the appearance of monitors

was most probable.

In

order to determine age, we amputated the last phalanx

from one of the fingers of a number of the monitor liz-

The specimen was processed by E M. Smirina(Moscow) to whom we owe our sincere gratitude.
The region of operations was regularly inspected and
ards.

all

points at which a tracks of the marked animals wasmeter.

The monitors were, however, distributed overthe space unevenly:

areas of high concentration alternating with thinly populated areas. Areas in which

populations were more densely concentrated (settlements) measure nearly 100-150 hectares. The distance between the centers of neighboring settlementsfrom 3-5 kilometers.
Each settlement isformed from a group of adult settled individuals conranges

sisting of five to six males of various ages and three toapparent were plotted on a chart. As a result, we haveat our disposal data on age, the location of homefour females.

ranges, and the nature of the interrelations between
the majority of the mature individuals settled in theinregion of operations.
Indirect exchange of information predominates inthepopulation(Tsellarius

and Men'shikov,

1994).

The monitor lizards comparatively rarely enter into
direct contacts.

A possibility of observations of con-tacts is all the more rare.

Therefore, in order to gather

sufficient data, we provoked contacts.

To this end. theIn each of two settlements which were

under constant surveillance, only one female took part
reproduction.

The rest of the females were notimpregnated over the entire period of observations
although they were in fact courted by the males. The

home ranges of all of the animals in a settlement
almost fully overlap each other. In the sparsely settled
areas between settlements,

home ranges may, to avarying degree, either overlap or be located at certain
distance from one another.

The home ranges of set-

1997

Asiatic Herpetological ResearchVol. 7, p. 110

tied animals range

from 30 to 200 hectares in size.

From time to time, each of the males makes brief, distant excursions beyond the borders of his home range,
during which time hemay visit the territory of aneighboring settlement.

It

is in

When the monitor examines any object

motion.

with its tongue (e.g. tracks of other animals, burrow
entrances and so forth), the lizard slightly lowers hishead.

should be kept in mind that

In ordinary conditions these behavior

and body

scheme. The real picture is com-of period
posture were observed during the most part

nomadic individplicated by the presence of stray anduals among the settled ones, altering the make-up ofof monitor's activity when for example the hunting
monitor travels over familiar area. "Confident gait"

The area of a settlement

may be generally characterized as a pose of routine

this is a simplified

the settlements and so forth.

amounts

to akind of "public property" for settled

inhabitants of thisguarded.

settlement.

Itisnot specially

We did not observe any territorial behavior

The
between settlements or within them.
encountersandunrestofaggressiveness during
degreeeitheractivity.

In this time the animal is in a state of psycho-

logical comfort.
it

A conflict of motivations is absent or

is possible to realize through the behavioral patternseveral motivations simultaneously (e.g. the search forfood and a sexual partnerinthe course of searchwith acquaintances was. however, incontrovertibly
lower than during encounters with strangers (Tsellar-

external circumstances do not pose obstacles to theius and Men'shikov 1994) and made the incorporation

fulfillment

newcomers

of

into

the

settlement

difficult.

The

behavior).

were under constant surveillance.
the region of research theInhibernation lastsof theexs 1. 7; fig 14a).
ples of behavior of monitors...",
This gait probably testifies to the self-confidence andemergence from hibernation, monitors widely travel
within their home ranges and intensively forage. In

threat.

is highperiod the social activity of monitors
with
into
contacts
enter
conspeenough: they readilycifics and show great interest in their tracks, but males

do not make any attempts to court the females in thisMatingperiod (Tsellarius and Men'shikov 1994).
it starts from the
a
short
time,
continues
forperiodfirst days of June and comes to end till the twentiethof June. Females, which have taken part in mating,or beginning of
dig the nest burrows in the end of Junetheclutchandduring 1-3
diligently protect
July,months after deposition (Tsellarius and Men'shikov
1995).

Socialactivityof otheranimals

rapidly

decreases during 1-2 weeks after the end of matingof moniperiod. Mobility and activity of the foraging
tors (females with clutches excluded) also decrease.
In the end of July the range, daily inspected by a mon-a third or quarter of daily range in May. In
addition, in the second part of the summer lizards
itor,

is

may, from time to time, spend several days in burrows
without emerging to the surface.

of activity

The "confident gait" was often observed in course

calm of the animal.

this

type

its

of encounters between monitors (see section "Exam-approximately from the beginning of October
During 1.5-2.0 months after
beginning of April.
to theof the animal andcorresponding(Ovsyanikov and Badridze, 1989).

weremajority of the animals mentioned in this article
members of one of two adjacent settlements which

The

physical state

prets

it

His contactant evidently

inter-

as a threat of attack or the possibility of such
In all directly observed instances the approachprovoked an unambiguous
gait"
The
approached animal manifests anxiety
response.
by"confident(see exs 7, 9 and Fig. 14, 16) or displays a peaceableness/submission (ex 1 ). The latter instances are rare.

Attack. When a monitor attacks a large prey he either

immediately rushes on it, abruptly starting a sprint, or
sneaks up to it before rushing from a distance of 3-5
m away. The sneaking up monitor walks with a rapid
step, creeping along the ground,often uses shrubs andrelief of the land as a cover. The act of lowering him-

self to the ground is also characteristic of the monitor

that is avoiding danger (Fig. 3b).

When sneaking up

and when rushing the monitor holds his head and tailDuring an ordinary hunt all observed
actions were directly connected with prey-catching.

horizontally.

Any special displays were absent.
of body of a monitor in assault

isIn general, posture

the same as in "con-fident gait".

Attacks on conspecifics were observed only in two
situations: a) when a female protects her nest burrow(Tsellarius and Men'shikov 1995); b) when an animal

The Most Ordinary Behavioral Acts of Desert

with a lower social status rudely breaks the "rules ofMonitorfor example, at
etiquette", suddenly appearing,"Confident gait". The animal moves calmly, carryThe monitor
ing his body high over the earth (Fig. 2).
lizard holds his head and tail horizontally and some-times the tip of the tail draws upon the earth while hea dis-

tance closer than individual distance (ex 4 and Fig.
12). In such circumstances the emotional state of theas anger or
attacking monitor may be characterized
are
which
expressed through sperage. Any signals
cialized posturesand movements during an attack

1997

Asiatic Herpetological Researchwere not recorded.

In

1992 we observed how a large

in a given settlement, andslightly lowering herself to the ground, and lunged at
him from a distance of 2-5 meters away. Biteress's

stealing up to his prey.

In all observed instances attacked animal

immediately

resorted to flight.

The "threatening gait". The

lizardmoves

direction of an adversary at a slow pace.

"sits down", liftingand looking around (Fig.
Very often animal assumes this pose when at great

6).

distance away is some disturbing object (observer, forexample).

"Sitting dog" posture probably

orientation response.

This reactionisisa classic

formed withpresence of weak fear (Hinde, 1970).

A monitor often raises his head during contact

behavior was, down to the most minute details, similar to the behavior of a monitor

age.

forepart of torso and head

had named Biteress and attempted to dig up the bur-

The female, having found the arrival at work, drew

I

"Sitting dog" posture. Monitor

few days to the nesting burrow of the female that weclose to him with quick dashes from shrub to shrub,IIrather closely linked to size, the duration of residence

monitor, considerably larger than the proprietress ofthe burrow, returned repeatedly over the course of arow in her absence (Tsellarius and Men'shikov 1995).

Vol. 7. p.

when the latter assumes
submission posture (exs.l, 3, 5 and Fig. 9). Thereforeit
may be possible that the "sitting dog" posture (Fig.
with subordinate individual6) or very similar pose also has a trace of dominance

the

in

His tail may

not touch the ground but may, on certain occasions,leave distinct imprints on both sides of his trail

in it.

"Sitting dog" posture was observed in course of

conflict between males,

when both contactants mani-

fested an unwillingness to yield (ex 9).

(giv-

ing the impression that the lizard is whipping his tailenroute).

When walking the monitor "drags his feet",leaving distinct tracks in the sand from the dragging
of his ringers and claws. We cansay nothing moredefinite about this pose because it has been noted
only"Stooping"". The

lizardslightly

lowers his head,

simultaneously inflating his throat and pressing his
tail to the sand(Fig. 15a). This takes place during theencounter of an animal with an unknown object in a
familiar area (e.g. a backpack, a shirt

hung on a bush),encounters whose course has been reestablishedin the event of a careless movement of the observer infrom tracks. The track of "threatening gait", however,hiding when the monitor lizard is not able to precisely

in

The female
very distinct from all other tracks.
leaves such a trail when she moves
away from her
isidentify the character and source of the movement, or

during a direct encounter with another monitor that is

nesting burrow in the direction of an approaching con-unknown or whose intentions the animal is not in a

Such trails have also been noted a few times

position to determine, i.e. during an encounter with an

specific.

on the part of the female who directed it at a male that

irritant that attests not to an obvious

pursuing her. The "threatening gait" was recorded
in encounters between males when individual distancepossibility of its beginning.

was abruptly broken (ex 4. Fig.

degree of fear, unease, lack of self-confidence. This
state can even be
unmistakably recognized in a liz-istion the threat

12).

In

such a situa-

was always displayed by animal withhigher "social standing"1.

In every instance, the ani-

mal to which the threat was addressed
sharplychanged the direction of his movement and in the
majority of cases moved away from the site of events,
sometimes taking flight. The lizards did not engage indirect contact.

statedanger but to the

In this circumstance theof the animal may be characterized as a weak

ard's tracks because his tail in these instances leaves a
distinct, straight

furrow in the sand (Tsellarius andMen'shikov 1994).
Zatir. If the uneasetion, theiscombined with strong excita-

monitor presses his cloaca and the hind partof his abdomen to the sand and crawls, leaving behind"Threatening gait" and attack were recordedinvery similar circumstances, sometimes "threateninggait" was followed by an attack. The state of the ani-

mal in "threatening gait" may beinterpreted as angera stripe of flattened sand (Fig. 5). This
stripe serves asa signal

olfactory

mark which combines incues(Tsellarius

anditself visual

Men'shikov

and1994).

Formerly considering the marking behavior of moniand Men'shikov
1994) we termed this act as "dragging" and marktors in a special article (Tsellarius

itself as the "drag".

When speaking of the social status of an individual, we hadin mind the"frequency of dominance". In other words, themore the monitors of a given settlementa subordi-

occupynate position in relation to a

The term is an unfortunate one as

being used for designation of another type ofactivity (Carpenter and Ferguson 1977). We propose
itis

to use the transliteration of the russian term "zatir".

given individual, the higher his

status. As one would
expect, the social status of males
turned out to be rather closely linked to size, the duration ofresidence in a given settlement, andage.

"Showing of the back". A monitor flattens the torso
dorsoventrally and incline laterally in the direction of

opponent, showing the back, as it were (Fig. 11. 13d).

At this time the tail is lowered, but not pressed, to theground and the throat may be slightly inflated. The
monitor always orients himself laterally towards a
threat.

1997

Asiatic Herpetological ResearchVol. 7, p. 112

On certain occasions one may note a tendency

ing the displays of passive intimidation. In the rarecases an infuriated monitor may fall into attack. The
state of the animal during "arching the back" is proba-

bly very similar to it in "showing the back" but thefear changes into anger
degree of fear is more high.

Aof "arching of the back".

We observed "showing of the back" when themonitor encountered retaliatory aggression (active

and rage as a degree of threat increases.

self-defense) on the part of prey and during the hesi-

the monitor will display extreme readiness for self-

behavior of a human who neither attempted todefense: he opens his mouth wide (Fig. 8a) and lunges

tantcapture the monitor nor made any sharp movements
and yet did not leave the lizard alone.
During"(Jape". If the danger is very serious and unexpected,

in thedirection of his adversary.

play attests to

Evidently this disWhen aan extreme degree of fear.

"showing of the back" wasmonitor, which is sleeping near entrance of burrow, ismore often observed in situations when, on one hand,being suddenly caught, he often convulsively moves
his legs wide apart, and feverishly turns his head withintraspecific contacts theitmay be suggested that the behavior of the conspe-

cific excites apprehension of the monitor, on the otheropened mouthpresent activity is rather
The emostrong and monitor refuses to take to flight.

"strength" of heartbeat

hand the motivationfortional state in such situation

may be characterized as

alarm.

inall

directions.
is

At

this

time anoticeably increased.

The

general picture shows strong resemblance to displaysof intense fear in man and other mammals (Darwin.

1872;Deryabin. 1974).

dence and does not disrupt the contact, the showing

The "gape" may be followed by an attack. The
animal becomes enraged, the urge for flight is curbed,

back monitor may fall into active self-defence (ex 8).

the monitor chases his adversary and, if he is success-opponent does not manifest any symptoms offear and confidently goes on with approach, the monitor commonly takes to flight. We have never observed

ful in catching hold of the latter, can be very difficult

If another lizard both

does not have self-confi-

If thethat attack or appeasement

followed the "showing of

However, during contacts with human thethe back".

"showing of the back" may fall into the "arching of

The latter may be followed by attack.

the back".

thissary, the tail is raised for ablow, and the headisturned in the direction of the danger) and elements ofpassive intimidation (i.e. the animal tries to appear
larger than he actually is: the back is arched, the tho-

rax is expanded, the bodyisraisedupon erect front

legs, and the throat is inflated) are combined.

Such posture is assumed by a monitor when thedanger is serious and paths to retreat are cut off.
"Arching of the back" is very common as a response
to a threat

on the part of human.

8b).

in

"Lurking". If during an encounter with danger thethat he has gone unnoticed, he will

monitor thinks

conceal himself by lying down and pressing himself

Furthermore, dependlizard
will either remain
the
on
the
circumstances,ing

tightly to the ground (Fig. 3a).

posture (Fig. 7a) adisplay of readiness for active self-defense (i.e. themonitor orients himself laterally towards his adver-"Arching of the back". In"Gape" was observed only
encounters of monitors with human.
to deter (Fig.

In such contacts it

prone or exit stealthily (Fig. 3b).
Itisdifficult to define themonitor in this situation.

emotional

state

of a

Some degree of fear takes

place for certain. However, "lurking" is not an involuntary display of fear unlike such reactions as "arching of the back" or "gape".

Itshould be kept in mind

that absolutely identical posture is typical of the stateof contentment (e.g. basking, resting of replete monitor in the

shadow of bush and so forth).

This thingshould be taken into account when the behavior of amonitor in intraspecific contacts is being interpreted.
It
may be important for interpretation that in an

sometimes follows the "showing of the back". We
have never seen this posture in encounters between

encounter with danger the monitor's eyes remained
open (in every instance of this that we were able toconspecifics.

make out) and in state of contentment the lizard very

If thehuman does not manifest hostile intentions,

often closes his eyes.

In the opposite

The "lurking" is customary in the social interac-case the monitor deals a blow with his tail and, after

tions of various species of lizards (Carpenter and Fer-given) or starts

guson, 1977) and has traditionally been interpreted
Such behavior is a usual
a display of submission.

the animal slowly retreats (Fig. 7b).
this, either takes flight (if a chance istomake a lunge in the direction of the aggressor.

Whenlunging the monitor turns to his adversary
breaking the lateral orientation and strongly decreas-ascharacteristic of the desert monitor also.

The female

often assumes this posture during the approach of an

I

adult male (ex

he

a

1

In this case the "lurking" may truly).

of submission,

display

we have neverbutobserved such a posture on the part of the subordinate
inVol. 7. p. 113

Asiatic Herpetological Research997

encounters of males.

On the other hand, however,

we have observed how it is assumed by the obviously
larger and stronger animal in response to a female's ora smaller male's display of readiness for active self-

defense (exs 2, 7 and Fig. 10. 14). We have neverobserved that "lurking" follows display of fear, uneaseor alarm during intraspeciric contacts. The primarysignificance of this posture is probably not a displayof subordinance as such but a display of a peaceableness.

In general, recumbent postures aremost char-acteristic of contacts of a "friendly" type (see below).

Flight. In encounters of a desert monitor with humanor another large animal the monitor seldom falls into

moment.

Usually a flight follows"showing of the back", "arching of the back" or "lurkflight at the firsting". If a danger arises at a distance more than criticalone (the latter is from 4 to 8 m) the animal will lurk asa rule.

In opposite cases the animal usually manifests

a readiness for active self-defence.

In the case whenhuman does not show aggressive intents, the lizard

Behavior Characteristic of Certain Types ofInterrelations
Manifestations of dominance. We

observed

The flight is not accompanied by any

nent's sight.

adisplaysrelatively

Very often flight taken place without any previous symptoms of fear (exs 5, 6 and Fig. 13). No

howmattertheinteraction

would

turn,

in

every

observed incidence of "sniffing", fighting, and ritual

combat,flighton the part of one of the contactants

asymmetry of behavior

more anxiety

another and another's tracks.

animal as a subordinant.

to

in

all

One monitor

in

We

with

contacts

regarded thefirst

Another lizard displays aself-confidence and was regarded as a dominant.
In the caseof dominancetoms".

of monitors, the behavioral syndromeis

divided into two groups of "symp-

On one hand there are distinctive features of

behavior which are connected with the social status ofan animal and which are manifest

inmany types of

These features are displayed from the firstactivity.
moment of interaction during direct contact between

On other hand there is the "behavior of the

lizards.

which is displayed only in agonistic contacts
and only when the "correlation of forces" has been
victor"revealed.

ing

The idea was formed that behavior of monitor durcontact with conspecific is more strongly

impressed by the social status of the animal (i.e. general experience and history of interrelations with con-specifics

nature.

ableof two always (or in overwhelming majority of cases)

special displays (Fig. 4).

During encounters between monitors in the majority of cases, a flight is probably an action of ritual

stableencounters between certain individuals.

will slowly retreat either stealthily (Fig. 3b) or keep-

ing the posture of readiness for self-defence (Fig. 7b)
and falls into flight (if falls) only outside of oppo-

wereclosely observe the interrelations of some monitors
for a length of 3-4 years.
During these years we

at

all)

than by concrete dominance andsubordinance. The lizard with low status even during

an encounter with his subordinant sometimes displays
the most anxiety than high-ranking animal in contact

with his dominant.

On the whole, the behavior of ahigh-ranking animal is distinguished by self-confidence: the monitor rarely displays signs of unease or.
rarer still, alarm.

This is especially apparent in track-

inevitably terminated the contact regardless of thestage to which contact may have progressed. In theing studies (Tsellarius and Men'shikov 1994). Absolutely dominant in one of the settlements, the fourteen

event of flight, the "conquered" generally retreated toyear old male named Vasya, boasting a snout- ventand a weight of 3.5 kg, didlength of nearly 600

a distance of 2-3 meters away,

more

rarely10-15

meters, resuming his usual pace afterward, with the"victor" generally not displaying intention to pursue

Exceptions to this rule are thosewhich one of the contactants immediately(exs 3, 5, 6, 8, 9).

cases

in

sets about an attack (ex 4 and Fig. 12).

well, however, the attacker's rage

In this case as

and the retreatee's

fear are certain only in the first moment of encounter.

From that point on both flight and pursuit begin to
increasingly take on the character of display (Fig. 12
It is symptomatic of this behavior that inand ex 4).

mm

not display substantial alarm even upon encountering
In Vasya's case we did not observe posturesof active self-defence (Fig. 7a) at all. As a rule, thea human.

monitor turned sideways towards the human encountering him, lowered his head, and slightly inflated his

The "showing ofthroat.

the

back"

faintly

away only at a slightly quickened pace, periodicallysitting and looking back (Fig. 6).

not one of the observed incidences did the attackerItcatch up to his adversary.

wasexpressed if expressed at all. In such a position themonitor first drew back a few steps and then movedishighlyprobablethat,duringencounters

between animals, behavior attesting to states of comfort are devoid of any expression of a conflict of moti-

vations (exs1.

7 and Fig. 14) and is in itself a display

Vol. 7. p.

1

14of high rank.

Asiatic Herpetological Research

We have observed such behavior prima-

males

in

which

have

high social standing.
Females may assume a submissive posture (ex 1 dur-

rily

)

1997

it.
For example. Mafiozi repeatedly visited the area of
Docentess's nesting burrow and even spent the night

with

her

in

a

singleburrow

and(Tsellarius

ing such an approach by a male but low-ranking males

Men'shikov 1995).

display a great degree of alarm and often resort totacts between these animals were not observed either

flight.

The specific "threatening gait" and lifting of

the head are also apparently linked to high social status.

Observers of monitor behavior have described anassumed when

"victor'spose" that isthe victorious monitor mounts theunambiguous

absolutely

one

It

must be added that sexual con-that year or later.

Characteristic of friendly contacts are the absenceor only very faint display of signs of unease on either
individual's part and the mutual display of "submission" (exs 6. 10 and Fig. 17). During such contact the
lizards never hold their bodies high over the

ground,1981a; Deraniyagala.
(Auffenberg,
1958; Gaulke, 1989; Horn et al.. 1994). A posture ofmuch as in normal movement or during the "threatenThe monitors lie down and either draw
ing gait".

this sort (e.g. topping, riding,

together with short, a few steps at a time, crossings or
crawl across in each other's direction. Expressions ofdefeated

istic ofstraddling) is character-many species of lacertilias and is displayed in

the course of both agonistic and sexual interactiona peaceful nature do not, however, impede the(Auffenberg, 1981a; 1983; Carpenter and Ferguson.
1977; Horn et al.. 1994; Noble and Bradley, 1933). In

sition of ritual combat (exs 5. 6 and Fig. 13).

the case of the desert monitor, we observed such pos-Common Ceremonies and Theirturing only during mating attempts but it is impossible

Function

to rule out its use as a "victor's pose" as well.

We designate as a ceremony those interdependent

propo-

Probable

one of the contactants assumes the posture of submis-

actions which occur during contact between two or
more individuals and are directed at the maintenancesion, the act of licking the "defeated" plays the role ofor establishment of certain social relations, and alsoa "victor's pose" (exthe rules

Within a certain context, it is possible that, when1).

In opinion of Auffenberg(1981a), tongue licking in the case of Yaranus benga-by which these actions are guided.

The

social status of the contactants, the dynamics of theirmotivational and emotional states, and the displays

lensis has a signaling function.

Displays of amicability. Informal contacts, founded

upon personal attachments, are common to many species of animals and may play a substantial role in theformation of the social structure (Panov. 1983b).

Aconnected with them determine the course of the cere-mony and its result for each of the participants but are
not linked to the essence (goal) of the ceremony itself.

Mutual "sniffing". Mutualis"sniffing"an almostsimilar sort of connection probably exists in the caseobligatory act in the encounter of two animals over aof Varanus komodensis (Auffenberg. 1981b),ben-

definite period of time. Exceptions may include those1983), V. rosenbergi (Green andcontacts connected with the protection of a nesting
burrow and cases of sudden, involuntary violations ofgalensis (Auffenberg.

King 1993) and some other species.

V.

In the caseofmonitors, the basis for this connection probably lies inindividual distance. In these situations one of the con-the habit of neighborhood as a familiar lizard evokes

tactants

much less unease than a strange one (Tsellarius andomitting the

Men'shikov, 1994).

It

may be possible that the per-spring,may immediately resort to threat or attack,"sniffing"

immediately

procedure.

following the

Only

in

early

emergence from

Mertens (1946) long ago noted the clearly expressedhibernation, and in the period when all regular activity
ceases before hibernation, was the "sniffing" proce-individual differences in character

among monitors.

dure rarely observed in encounters between animals.

the case of theMonitors encountering one another often (albeit notsonality traits of an animal are of significance as well.

Accordingtoour observations,indesert monitor, individual differences in the degree of
excitability,

aggressiveness, and thealways) did not engage in contact at all.

ability to alter

behavioral patterns in changed circumstances may bevery pronounced.

We only observed friendly contacts between settled monitors whose home ranges werebroadly over-lapping for a long period of time. The aggressive
reaction of animals during such contact may be suppressed to such an extent that the female guarding her
nesting burrow will allow another individual to visit

During "sniffing" the monitors usually

first

lick

snout of their conspecific. then his side, thesacrum region, and the base of the tail. Sometimes
the"sniffing" proceeds without displays of alarm or
unease on the part of the contactants for the durationof the ceremony (exs 6, 10 and Fig.
quently,

17).

More fre-however, unease or alarm occurs.

Itisextremely common for animals to display alarm after
mutual "sniffing" of the snout and to try to avoid

Asiatic Herpetological Research

1997

being licked in the sacrum region.

In these instances.

the monitors "waltz": they circle, as

if

attempting tocatch up with each other's tail (Fig. 13d. 15b).

both of the animals always "shows the back".

One orIt

theVol. 7. p. 115

always the one which originally manifested alarm to
the greater degree, dealt a blow with his tail whichinvariably put his adversary to (light. The larger male
did not necessarily emerge victorious from the fight.

contactants begin to display alarm at the very start of

Nor was the victor necessarily a resident of the settle-

contact, then they may immediately orient themselves

ment on whose territory the encounter took place.

not to the facial but to the sacrum region, which inevi-

The frequency of fights probably depends on anumber of circumstances but first of all on the social

tably leads to "waltzing".

The motivation for "sniffing" is. apparently, quite
Animals frequently draw together and do notstrong.

interrupt their contact until each contactant has licked

structure of the population in a given area and at agiven time. In other words, it depends on the predominance of a certain type of interrelation. The social

the other, even when the other evokes in each a strong

structureunease (ex 8).

ture.

The primary goal of "sniffing"is probably theofcertaininformation
about
a conspecific.
receipt
Familiar animals are probably capable of recognizing

each other by some external features.
In every
instance of contact between unacquainted individuals,

isnot an unchanging, species-specific fea-

Although the interrelations between monitors in
the region of studies from 1991 to 1993 may be characterized as highly peaceful, in the previous period
these relations were, evidentally. of a rather different

The majority of monitors caught for marking
1990, males and females alike, had fresh, deep

nature.
inorwoundshowever, sex and reproductive state were determinedscars

only by olfactory means (Tsellarius and Men'shikov

region or on the shoulders, rarely on the side.

1994).

Therefore, it may be possible that the urge to

one's conspecific has as

lick

its

basis the urge to

receive information about the physiological status of
the encountered individual.

locatedprimarilyinthesacrumThesewounds are reminiscent of the teeth tracks
which are left when a monitor takes something in his"mortal grip". In 1991 wounds and fresh scars were ascars andrarity,'and from 1992-1993 were noted two times in

all.

from

Apart

this,

to

however,mutual"sniffing"

be a required ceremonyin theappears thereby
course of which the social status and the personal
interrelations of the contactants are determined (orRitual combat. Ritual combat is that type of agonistic

interaction in which animals enter into direct con-tact butmeasures which could lead to the mutilation

The analogous significance of the cere-of an opponent are excluded. Until now. ritual combat

mony of "sniffing" has been well known for socialized
species of carnivorous mammals, canids in particularin the case of the desert monitor has gone undocumented, although it has been described for many other(Schenkel. 1947: Lorenz 1969).

species of this genus (Green and King 1993: Greer.

confirmed).

Fight. We designate as a fight that type of agonisticinteraction in which: a) monitors enter into direct contact, b) measures are taken that can lead to the mutila-

tion or deathof the contactants. c) on both sides

displays of anxiety and readiness for self-defence take

place

in

the course of the interaction.

Following
Auffenberg (1981a) and other researchers, we treat
blows of the tail as measures which can cause mutilation, although, strictly speaking, in the case of moni-

tors they are a ritualizedmeasure, a lesser one.

incontrast to bites which are likely to inflict injury on an

In every instance, ritual combat is

we observed three instances which may besome stretching the
In each of the three instances unknown or

qualify as fights, although withpoint a bit.

preceded by theceremony of "sniffing" (exs 5, 6 and Fig. 13). Combat
may be broken off at the initiative of one of the participants at any moment and in the majority of cases thistook place at the very first stage of combat, the "cross-ing of necks" (ex 6). We only observed one instanceof all-out ritual combat (Fig. 13). Every instance of
ritualcombat or the attempt to propose it was noted

only during contacts between animals from the same
settlement.

combat was recorded only in the case of

Ritualopponent.
In all,1989; Horn et al. 1994).

males.

It

may be possible, however, that such combat

also takes place among females.

At the end of June

1993. we observed the tracks of an encounter between

unfamiliar males from various settlements entered

two females from the same settlement, Frosya, five

Displays of alarm and unease wereclearly expressed by both sides (ex 8, Fig. 15). Fol-

years old. and The Fourth, four years old. The trampled patch of sand that remained at the sight of theirlowing the "sniffing", accompanied by the "showingof the back" and "waltzing", one of the animals.

encounter was somewhat similar to those that are left

into contact.

combat between males. Frosya followed
The Fourth's tracks for nearly 45 m to this patch. The
after ritualfemales were obviously within each other's field ofvision and both, probably, "showed their back" from

From the trampled patch the females
headed in different directions. Notably. The Fourth

ritualm and Frosya moved at an easy pace.

behavior connected primarily with the displayof dominance and is not a ritualization of the fight.

time to time.

ran for nearly 17

1997

Asiatic Herpetological ResearchVol. 7. p. 116

In the caseof the desert monitor and some othercombat is extremely rare in
The reason for this probably lies

varanids. all-out ritualnatural conditions.

In the case of certain varanids. ritual combat has beenin the fact that a certain combination of circumstances

noted between males, between females, and in contacts between the sexes (Auffenberg,1981a; Gaulke.

must be present for ritual combat to occur: unceranitainty should exist in the interrelations between1989).

mals, their social status should be about equal, anddominance in relation to one another unestablished.

The significance of ritual combat for varanids is

In so doing, in the caseof an obvious inequality

inunclear (Greer, 1989; Horn et al.. 1994) and attempts
to link this type of interaction with the struggle for ahave
specific resource (e.g. food, territory, females)

dominance is resolved by the process of "sniffing" andnot been successful. In those cases where a few males

direct struggle to establish seniority is unnecessary.

competing on account of a female, skirmishes
have been observed but ritual combat did not takeof foodplace (Carter, 1990). Encounters near sources

strength (i.e. in thearemay lead to a form of ritual combat but more often

majority of cases) the question of

the conditions of a stable settlement, socialIn

rearrangements, taking into account the long life
expectancy of monitors, are relatively rare: the ani-

Gaulke.

mals know one another personally and the rank ofeach is known to all the rest. The need not only for

majority of varanids,

combat but even for the sharp display of dominanceincluding the desert monitor, has not been established(Green and King, 1993; Greer, 1989; Tsellarius et al..

destabilized to the point that a large number of lizards

proceedotherwise1989).

Territoriality(Auffenberg.
for theIn the case of the desert

1994).

1991; Tsellarius,

1981b;

rarely arises.

appearIn the event that the social structure isthat are not sufficiently wellknown to one

monitor it is precisely these skirmishes which, stretch-another, the social rank of an animal is more likely to

ing the point a bit, may be regarded as territorial (i.e.

be established in a series of fights and the conditions

the encounter of a settled male with an unknown ani-

for ritual combat rarely take shape.

mal, a female's guarding of her nesting burrow) proceeding more in the form of a fight than ritualized

Examplesinteraction.
If

we examine

ritualcombat

in

the

case

of

made

varanids on the whole, then the impressionthat the essence of combat consists in the mutualisattempt to knock over the opponent and assume the"victor's pose".

The animal throws his front paw (orof Behavior of Monitors During
Encounters with Conspecifics.

The encounter of a settled male with a female froma neighboring settlement. May 26, 1993. The ContacI.

tants:

the male Mafiozi, six years old,

SVL 475 mm,weight 1.3 kg; the female The Fourth", four years old,
SVL 425 mm, weight 1.2 kg. Over the course of two

the front and the hind) over the back of his adversary

years Mafiozi had encountered The Fourth's

and tries to stand up over him. This is quite apparent
in both our photographs (Fig. 13) and in detailed

until this moment probably had had no direct contact
with her, in this year in any case. The Fourth had

different

species of

Davis

al.,

been released the previous evening into a burrow
located seven meters from Mafiozi's shelter that night

descriptions ofvaranids

combat among
1981a;(Auffenberg,

et

1986;

Deraniyagala, 1958: Gaulke, 1989; Hom et al.. 1994).
This grappling, the attempts of each opponent toattain the "victor's pose"

while simultaneously trying

to frustrate the other,

the essence of the combat.

isThe struggle in a standing position on the hind legsa moreevidentally developed as an attempt to occupy
favorable position for toppling an adversary.

It is

sig-

end has been
finished by "victor's pose" (Deraniyagala. 1958: Hornnificant that a struggle attained to the

et al.,

1994).

If this is true,

then ritual combat is most closely

tracks but

and in the morning came to the surface later than hedid.

On the morning Mafiozi, lying near his burrow,without hesitaspied the walking female, rose, and,The femaleforher.
headedofunease,tion or displays

immediately lay down and Mafiozi. having reachedher, licked her head and sacrum region. After this hemoved off a few steps to the side, raised his head (Fig.

and lay down, turning away from The Fourth.
Mafiozi twice more approached the female, with aninterval of a few minutes between, and again licked

9)

her.

Only after this did he move away.

When thelinked not with the fight for a specific object but withpurely social interaction.

Combat is a development of2

A11 females mentioned in this section of article did not takepart in reproduction for

all period of investigations.

male had left. The Fourth, not rising, lifted her headbut lowered it again as soon as Mafiozi turned towards
After every lick of his tongue, Mafiozi lifted hisher.

head high and held it this way for some time.

The encounter of an adult, nomadic male with a2.

Vol. 7. p. 117

Asiatic Herpetological Research

1997

The animals aresame settlement and knew one

the males in the given settlement.

members oftheanother well for not less than a year before theencounter. None of the conflicts between them before
the one described below, or for a year and a half fol-young, settled female. May 14. 1991. The course of
events was reestablished from tracks. The Contac-

lowing it, have been noted. On the morning, Mafiozi,
walking around a bush, literally stepped on Feodor.

SVL

who was basking in the sun after emerging from histants: the male Grigory, seven to eight years old,mm, weight 1.9 kg; the female Frosya, three
Grigory
years old, SVL 425 mm, weight 1.6 kg.
505

summer appears

hibernates in this region and in thehere episodically, one or two times a month.

Frosyanight shelter (Fig.

ma shrub from different sides (Fig. 10). Judging by the5direction. Frosya also lay

down.

After

some time

Frosya rose and. walking around the male, left in thesame direction she had come before the encounter.

Grigory followed her tracks for nearly 50 meters,leaving powerful "zatirs", and then also moved in the
direction he had kept to before the encounter.
3.

The encounter of an adult, hut never impregnated,settled female with a settled male. May 22, 1993.

The

Contactants: the male Es the Ninth, eight years old,

SVL 480 mm, weight 2 kg; the female Frosya, five
years old, SVL 450 mm, weight 1.2 kg. The animals
have belonged to the same settlement for no less thanfour years and are well acquainted with one another.
In 1992, Es the Ninth persistently, but unsuccessfully,

courted Frosya.

Frosya encountered Es the Ninth onmthe length of their step decreased. Feodor
then shifted to a walk and followed Mafiozi for about

next 15

Frosya immediately turned sideways to themale and. probably, "showed her back". Grigory lay
down at once. Having taken a few short steps in his

The latter jumped on him.

warning, and Mafiozi jumped
apparently
first 15took
toaside andflight. Evidentally. for the
For theas
could.
ran
as
fast
the
monitors20
they

has lived in this area permanently for at least two
years. The monitors came into contact while circling

tracks,12).

without

m by the "threatening gait". As soon as the young
male also shifted to a walk, however, Feodor made aof speed, compelling Mafiozi to again take
The chase continued in this fashion for more
flight.

burst

than 200 m.

Finally, having startled Mafiozi again inroutine order, Feodor shifted to an easy pace and.
sharply changing direction, went to one of the nearestcolonies of Rhombomys opimus where he began tohunt. Events proceed in like manner when a female ischasing a monitor who has encroached upon her nest-

ing burrow.
5.

Ritual combat between two settled males. June

1993.

The Contactants:

1,

Mafiozi and Edik. both 6

years old, measure SVL 475 mm, and weigh 1.3 kg.
The animals know each other well and have been
members of the same settlement for at least four
years.

At the end of 1992, the leader of the settle-

ment, the old male nicknamed Feodor, disappeared.
Among the remaining males Mafiozi is one of the

He is active over theand most energetic.

morning when he was laying, half of his bodysticking out of his burrow, and slowly headed for him,

entire space of the settlement

lowering her head and slightly inflating her throat. Es

beyond its borders. Edik's primary region of activity
lays on the northeastern edge of this settlement and hethethe Ninth, obviously disturbed, made a movement in

her direction, stooping slightly.

Frosya "showed herback" and began to move off sideways, not letting themale draw right up to her (Fig.

1

1 ).

Then she ran a

few meters away and the monitors froze. After a few
seconds of immobility, the female slowly moved
away, frequently looking back. Es the Ninth remainedin place until

she was hidden from view, watching herfrom behind in theattentively"sitting

dog" posture.

licked her track,left

a zatir,

and regularly goes far

The encounter
The monitors spent thevisits its central area only episodically.

took place on the morning.

night in the same colony of Rhombomys opimus andemerged from their burrows almost simultaneously.

Ithighly probable that the animals had made contact
in the burrow. The lizards lay for a few minutes nearisthe exit

from the burrows and then Mafiozi moved

Then hetoward Edik who, in his turn, took a few steps in his

and also

direction.

moved away.
4.

largestA skirmish of two settled males that resulted fromthe violation of individual distance. June 26.

1991.

The course of events was reestablished from tracks.
The Contactants; Feodor, nearly 10 years old, SVL
550 mm. weight 3.0 kg; Mafiozi, 4 years old, SVL
430 mm, 1.5 kg. Feodor has the highest status among

Meeting, the monitors lay down and, lying

down, licked one another first on the snout and thenon the side and sacrum (Fig. 13a). Next, they crossed
their necks, each attempting to deflect the neck of his

opponent to the side while simultaneously trying toseize with his paw the supporting foreleg of rival (Fig.
13b).

Until this moment neither of the monitors had

displayed any alarm or substantial unease.

Then.

1997

Asiatic Herpetological ResearchVol. 7, p. 118

Mafiozi succeeded in budging his opponent (Fig. 13c)and the monitors began to "waltz" (Fig. 13d) at whichtime Edik (left) "showed his back", displaying obvi-

himself, ran a few meters off, switched to an easy
and went off
pace, and left. Mafiozi licked his tracks
in a different direction.

At some moment the monitors were side

7. The encounter between a settled male and a nomad.

to side and their heads were directed in the same way.

Mafiozi immediately threw his front paw over Edik's
shoulders. The latter did the exact same thing at once.

May 24, 1993. The Contactants: Alitet, 5 years old.
SVL 470 mm. weight 1.4 kg; Shot Glass. 10-1 years
old. SVL 560 mm, weight 1.8 kg. Alitet is a settled

The lizards made attempts to overturn each other, slip-

residentous alarm.

One

neck.
ping their heads under their opponent's

1of thesettlementon whoseterritorythe

Shot Glas has spent the winterencounter took place.

time that Mafiozi succeeded in doing this, the moni-

within the borders of this settlement for at least three

tors rolled over on their backs and ended up in theirshifted to theprevious position. The lizards graduallyof hibernayears but about a month after coming out
tion goes far east and appears here only episodically,"face to face" position (Fig. 13e) and. continuing thefor two to three days, until the end of the season ofontheir hindlegs.

Havingassumed a vertical position and closed their front
paws on each other's backs (Fig. 130. the males construggle,

began

to

rise

tinued their vigorous attempts to topple their oppoThe males fell down together a few times,
nent.

breaking their grip

the

in

process (Fig.

but

13g),

immediately jumped up and seized one anotheranew. The entire skirmish was conducted very enerof the opponents changed
getically and the positions
andran
to the side for 3-4
rapidly. Edik suddenlymthe monitors watched one another motionlessly for

moved away,nearly 30 seconds. Edik then slowly
andfrequently looking back.
describing a zigzag
Mafiozi,

remaininginplace,

him

watchedfrombehind, raising his head. The entire struggle, from themomenttheadversaries

longer than two minutes.

crossednecks,

lastedno

The entire encounter, fromthe moment the monitors emerged from their burrows,
lasted 14 minutes.
6.

The Contactants:

Rhombik, 5years old, SVL 445 mm. weight 1.2 kg. Mafiozi
older and larger (see ex 5 for his description). TheisAs far as we know, these monitors had not

had direct contact this year, although they regularlycame across one another's tracks. The encounter took
place on the morning.

The monitors saw each otherof nearly 15
practically simultaneously, at a distancemeters, and froze motionless for some time. Then.
Shot Glass decisively headed toward Alitet (Fig. 14a)

who. having allowed him to approach within .5-2.0
m. inflated his throat and "showed his back". Shot
1

Glass immediately lay

down (Fig.

14b) and Alitet.

moni-

stepping slowly, walked around the recumbent
tor from the side and, looking back from time to time,

went away. About a minute later. Shot Glass rose and,to the ground, licked Alitet's tracks
14c). left a zatir, and followed the trail of hispressing his
(Fig.

tail

for nearly 34
conspecific. Having followed the tracks
m. he "sniffed" Alitet's excrement and peacefully

went away, sharply changed direction.
8.

Encounter and ritual combat between two settled

males. June 2, 1993.

activity.

An encounter between two unfamiliar males which

ended in a fight. May 31. 1993. The Contactants: The
1.4 kg;Tip. 6-7 years old, SVL 460 mm, weight
The
2.2
Tip
kg.
Dusty. 12 years old, 590 mm. weightisa settled resident of the settlement on whose terri-monitors belong to the same settlement and have
known each other well since at least 1991. On theat least three years.
tory the encounter took place. For
borders of this setthe
Dusty has spent the winter nearevening of June 1. 1993. without our interference, themonitors spent the night in the same colony of Rhom-

tlement but immediately after coming out of hibernation has traveled to places located several kilometersbomys opimus which that day. in the morning, ayoung female from a neighboring settlement. The
Fourth, had repeatedly visited. Emerging from theirto the southeast.

burrows the next morning, the monitors lay for a longtime near each other, basking in the sun and yawningInhibernation for the

1993 he remained in the region ofsummer for the first time.

encounter took place early in the day.

The

Having noticed

Their behavior was not unlike the usual

The Tip at a distance of about eight meters. Dusty lay
for a few seconds and then headed for The Tip who
immediately turned sideways to him. The Tip dis-behavior of monitor that have emerged from his night
shelters. After approximately 30 minutes the animals

played greater and greater alarm as Dusty approachedand the "showing of the back" became more and morecrawled together and the procedure of mutual "sniff-

inflated throat,
apparent (15a). Dusty's lowered head,
and tail pressed to the ground appear threatening butof self-confiactually attest to his unease and lack

in

turn.

ing" lasted for nearly 10 minutes.

It

was not accom-

panied by any apparent displays of unease. Mafiozi
then crossed necks with Rhombik and attempted to
freedpress him to the ground. Rhombik very calmly

dence. After the mutual "sniffing" of the snout. Dusty
but theattempted to "sniff" The Tip's sacrum

latterAsiatic Herpetological Research

1997

drew back, "showing his back"

(

15b).

As a result, themonitors described a few circles in place.

Suddenly,

The Tip dealt a blow of his tail and Dusty quickly

Vol. 7. p. 119

meters off, shifted to a walk and alter 25-30 m enteredof Rh. opimus. C-59 immediately assumed anormal position and. once The Tip was hidden froma burrowscurried aside. Having moved about six meters away,
he stopped and the monitors watched each other for

view, walked about the site of the encounter, inspect-some time. Then Dusty slowly moved away, barelylooking back. After one or two minutes The Tip also

10. A friendly encounter between an old male and ayoung male. May 12, 1993. The Contactants: Chuck
Norris, not less than 10 years old, SVL 520 mm,
weight 1.8 kg; Egghead, 4 years old, 425 mm, weight

left, not even having beeninterested by his opponent's

tracks.
9.

An encounter between two unfamiliar males, June

The Contactants: The Tip. 6-7 years old;
C-59, approximately the same age as The Tip but a bit
larger, his SVL is 480 mm and he weighs 1.5 kg. CIS.

1993.

59's

home range is situated in a zone little settled bying burrows, and then peacefully went away.

1.2 kg.

The home ranges of both monitors broadly

overlap but lay beyond the boundaries of the region
where regular observations were conducted and theirandstatusthehistoryof

theirinterrelationsisunknown.

monitors between two settlements located far from

Having caught sight of each other, bothmonitors lay down and for a long time crawled around

one another. Hence, he periodically makes excursions

each other, "sniffing" their partner's snout, side, andbeyond the borders of his range. The previous
evening C-59 was released into a burrow not far fromThe Tip's night shelter in almost the center of the lat-base of the

far

ter's

this.

C-59 was caught nearly a day before
Over the previous three and a half years he had

settlement.

appearedin thegiven region not more than two or

unease.

tail,displaying

off their activity and for 10-15 minutes

near tolie

each other, occasionally closing their eyes (Fig.
After this the "sniffing" was resumed.

17).

At the end of

the encounter the young male began to make "zatirs"

Emerging from his night shelter in theearly morning, C-59 walked for some time about theneck in doing so, after which he left.

colony of Rhombomys opimus and, finding the burrow
where The Tip had spent the night, hid in it. About 10

immediately entered a burrow.
minutes in all.

three times.

no signs ofvirtually

From time to time the animals would breakmarks), crawling across Chuck's back and
The old male(signalContact lasted for 47

minutes later the monitor again appeared on the surface from a different opening, having traveled nearly
three meters underground.

Emerging from the bur-row, C-59 looked around, licked the sand, yawned,
and. having moved a few meters away, lay under abush, head raised.

The Tip emerged from his own

burrow a few minutes later and headed confidently for

C-59 who remained in the very same posture but kept
an eye on The Tip, turning his head to follow the latmovements.
The Tip made a circle, having
drawn close to C-59 who at that moment rose halfter's

Discussion
On a certain stage of investigation of social structureof population the researcher will inevitably clash withnecessity of a studying of mechanisms of the social
reciprocal influences, that is the problem of intraspecific communication.

In classic ethology the concept of animal commu-

nication is based on the three postulates:
that

posed

behavioral

acts,

that

a)

carry

it

is

pro-

socially

important information, must be rather exotic, in orderway, inflated his throat, and "showed his back" (Fig.
16a).
Later. The Tip made yet another circle andthat a perceiving animal will be able to correctly sin-

Both monitors assumed thegle a communicative signal out of series of non com-again returned to C-59.

routinebehavioral"sitting dog" posture (Fig. 16b) and remained motion-municative,

less for some time, located at a distance of about two

communicative act must be very stereotyped, in order

meters from one another.

Then The Tip again began

to make a circle, at the same time inspecting the bur-that a

acts;

b)

a

monosemantic interpretation will be provided;c) a set of responses to certain act must be strictly lim-rows of Rh. opimus as if not noticing the newcomer.

ited to provide its adequacy and coordination of inter-

When he came near to C-59 the latter jumped up andThus, a communicative system is being
considered as the system of discrete, ritualized behav-

turnedhaving lowered his head,inflated his throat, and flattened his trunk dorsovenin

hisdirection,

trally. The Tip did the same and the monitors simultaneously lunged at one another. Each having struck thesacrum ofhis

opponent with

his

nose,

and each"showing his back" (Fig. 16c), they made a full revolution after

which The Tip jumped aside, ran 9-10

actions.

ioral actswith fixed significance (Hinde.1970; Mac-

Farland. 1985).

A process of communication is being

regarded as the stereotyped succession of acts, which
is founded on either innate or learned automatism of
responding.

1997

Asiatic Herpetological ResearchVol. 7, p. 120

These three principles are formulated by Tinber-them the most important ones are such interdependent

gen and his followers in course of investigations of
behavior of invertebrate animals, pisces and aves. It

factors as a) initial emotional state of contactants; b) a

should be noted that behavior connected with repro-

status of animal,

duction (courtship, mating, parental care) was mainlyintercourse with conspecifics.

history of interrelations of given animals; c) a social

examined. This behavior is directed at the reaching of

An investigationsof "everyday" social behavior (especially in mamlimited set of strictly specific aims.

mals) led to accumulation of facts which poorly har-monize with basic principles (for survey see Panov,1983a). Evidently, it should be admitted that development of communication on the base of other princior

ples

on

simultaneously

the
is

base

of

severalprinciples

i.e.

personal experience of previous

When a problem of decoding of any system of signals arises, it should be useful to formulate a supposition of what a kind of information
this system.

istransmitted by

It should be quite correct to use an analy-

of nonverbal communication of our own species
for solution of this problem. A majority of nonverbal
sisbehavioral signals of Man carry information about theemotional and/or motivational state of an individual.
Direct information about the intents of an individual

possible.

A contradistinction of exotic behavioral acts of theor his external circumstances is not contained in theand quite useless both for ascertaining of communica-

Nonverbal behavioral
majority of behavioral acts.
signals carry information about circumstances and/or

tive significance of these acts and for analysis of their

the intentions of given individual so far as a particular

In the point of view of aforming and origin.
researcher, who observes the monitors during all peri-

external circumstance frequently provokes a particular emotional state and this emotional state is. in aods of their activity, the "confident gait" is an element

particular situation, a base of a particular action.

of common, routine behavior, and the "showing of thehuman's response to behavioral signal may be very
diverse and depends upon the situation, his emotional

desert monitor and non-exotic ones

back"

is

a bright, exotic posture.

isvery relative

But

in

monitor's

point of view the "showing of the back" or "stooping"are far more usual acts than "confident gait", sincemonitor observed his conspecifics during contacts
only. Only the monitor's "point of view" is importantfor formation and evolution of his communicative sys-

A coincidence of researcher's and monitor's

tem.

"points" about exoticness of the posture should bepossible only in the case if a monitor would be awareof his behavior and identify his own behavior with
As far as we can judge bythat of a conspecific.

behavior of monitors during contacts, all features ofstate

and his personal experience.

It

is

A

evident that

communicative systems of other mammals is organized in a similar way (Schenkel. 1947; Ladygyna-

We have not any reasons

Kots, 1958; Lorenz, 1969).
to expectsome things of a fundamentally different

nature in other higher vertebrates.
isIt

strictly

imagined that the majority of behav-

ioral acts of the desert monitor clearly reflect the emo-

tionalstateof an animal and are

in

this

regardperfectly unambiguous. But displays of even diametbehavrically opposed states may contain identicalbehavior .exotic and non-exotic, are equally important

ioral elements.

for communication.

emotional state may be various and, accordingly, theEmotional
intensity of display may be various too.

A large majority of behavioral acts of desert monitors may

be expressed in varying degrees:

be very distinctive or hardly noticeable.

they may

Many differ-ent acts may smoothly turn one into another. Somesignaling elements of behavior, inflating of the throat
for example, take part in many displays,
distinctly different significance.

Itwhich have

was noticedin

In addition, thestates are not discrete and,intensity of every

between them, there exist
It concerns the mani-

an entire gamut of transitions.
festation of these states also.
It

seems to us that, without an understanding of

the dynamics of the motivational and emotional states

many species (Auffenberg,1981a; Panov and Zykova.

of animals engaged in social interaction, a correct
andinterpretation of the course of these interactions

1986; Hikida, 1989) and. evidently, it is not an excep-

their result is often complicated or altogether impossi-

tion, but a rule.

ble.

The data from our observations conforms poorly

A human perceives the behavior of a conspecificto the widespread perception of lizards as animals thatas a stream of integrated mental pictures.

display relatively primitive behavior based upon simple stereotypical reactions. A behavior of the monitor

elementary behavioral acts, as a rule, are not perceived individually. This stream of nonverbal signals

during encounter with conspecific is definitely not an
automatic response to the behavior of partner. Behav-

ceiver,ior

isconditioned bymany circumstances, among

first

Separateof all influence the emotional state of the perand do not act upon behavior immediately.

Adjustment of behavior is being realized by indirect

Asiatic Herpetological Research

1997

way through a change of subjective feeling of situation and emotional estimation of it.

ble that theItisquite proba-mechanism of the animal's perception ofVol. 7, p. 121,1968: Simonov. 1970). Probably emotions are a verymechanism of estimating the influence ofancient

internal

and external irritants, which developed longconspecific's behavior is the same. If this suggestionis right, we arrive at a picture of nonverbal communi-

before arising of new cortex. Morphological and neurophysiological ground for forming of emotions prob-

cation as a "process of tuning each of communicants

ably

A

long ago as the anamnia arose(MacLean, 1949; Dethier and Stellar 1967). A pres-mechanism of this tuning is influence of animals theence of afferent tone is the indispensable component

into the behavior of its partner" (Panov,emotional

andmotivational

state

of

1983a).

each

other

arose

as

of conditioning reaction, without it a forming of feed-through the exchange of information about the alteration of these states. Communication of this sort asks

back mechanism is impossible (Wiener. 1958).

for analogy to our own highly

developed systems ofnonverbal communication, such as music, dance, etc.

and mammals (MacLean, 1978; Regal, 1978; Tsellar-

During many decades amidst ethologists it wasconsidering as an indisputable tenet that animal's sub-interactions, universality

the majority of lizard's signals (Auffenberg,

jective feelings (including emotions) are unable to be

1981b; 1983; Carpenter and Ferguson 1977; Gaulk,

subject of scientific analysis. Actually it meant thatexistence of subjective feelings in animals is denied

for education (Brattstrom,

since a veto upon the use of this idea for explanationof mechanisms of behavior was imposed. But till now

no one succeeded

Essential resemblance of the behavior of lizards

and Men'shikov 1994), unstereotyped course ofand non-discrete nature ofius

1981a;

1989; Panov and Zykova, 1986), rather high ability
1978; Krushinsky, 1977)

make quite permissible the supposition that reptiles
have rather developed forms of psychological activity.

creating a general theory ofinbehavior within the bounds of behavioral approach.

andAcknowledgments

neurophysiologists, having
Psychologists
looked at the matter the other way round, propose a

The work was financed by the Uzbek Zoological Farmand the USSR State Committee for Nature Protection

concept of evolution of the psyche of vertebrate animals and influence of psyche on their behavior. This

and, following the disintegration of the Soviet Union,

concept well conforms to facts (Anokhin, 1968; Delgado, 1969; Shepherd, 1987; Vartanyan and Petrov,1989).
Using a concept of emotion, one generally

through contributions by the firm ALGA-ECO (Director,

S.

V. Tsvetkov)

and the Joint-Stock Company"The LIKO Firm" (President, S. V. Vlasov). and at theauthor's own expense.

The authors are deeply gratefulwork: M.

managed to obtain the harmonic, economical explana-to the volunteers who took part in the field

Suppositions were warily
declared that not only emotions, but also the higherV.

tion of observed behavior.

kinds of psychological processes, those are termed a"mind"

in respect toMan, are characteristic of ani-mals (Gallup, 1985; MacFarland, 1985; Sevastyanov,

The existence of emotions is not called to1989).

question in respect to mammals with a developed
brain. The analogy of basic and some secondary emotions of Man and other mammals

is

not refused also.

But in respect to reptiles, a use of the concept of emo-

Borovkova, Dr. V. A. Cherlin. S. G. Davtyan. Yu.

Derbyshev, A. Kuznetsov. N. Grefner, Dr. A. V. Gromov. T. Makarova, Yu. G. Men'shikov. K. Mil'to. A.
M. Murashev, A. Naryshkin, E. Yu. Portnov, Yu. Vostrukhina and Dr. A. M.Zakharov.
the completion of this researchpossible.

Without their help

would not have been

We would like to express our gratitude to

Dr. N. B. Ananjeva (St. Petersburg), Dr. 1. S. Darevsky(St.Petersburg) and Dr. A. D. Poyarkov (Moscow) for
I. Atamuradov (Ashkhabad). Dr.

their advice, Dr Kh.

Howwe denied the existence of a sharp border
between the psyche of Man and that of other mammals, we find ourselves before the necessity to seri-

T K. Kadyrov (Kyzylkum Preserve), Dr. S. A. Shepi-

ously warrant a placing of such boundary-line in any
other case.

processing the film, S. V. Chistyakova for her help in
preparing of manuscript of this article. Douglas

Emotions are an internal regulator of psychologi-

Greenfield and Betsy Lauppe (USA) for their help in

tion

is

unusual for the majority of zoologists.

ever, iflov (Tashkent), Dr. A. E. Subbotin (Moscow) andDr

O. I. Tsaruk (Tashkent) for their help in organizing thestudies. I. A. Mukhin and E. Petukhov for their help in

On behalf of all

cal activity and behavior, and are a universal measuretranslation of the article into English.

of values that have a great adaptive importance.
Under the shortage of prognostic information or

the participants in the fieldabsence of possibility of processing of it the emotional estimate of situation allows one to quickly findother local residents who provided fresh water, provi-

one's hearings and to

make one's choice (Anokhin

work, we would like tothank K. N. Jhankaraev, A. Annagel'dyev, and thesions, and mail for our camp every month even when
we were unable to pay for its transportation.

1997

Asiatic Herpetological ResearchVol. 7. p. 122

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Asiatic Herpetological ResearchVol. 7. p. 124

AppendixI

Figures 1-17

Figure 1.

Landscape of the region of operations, (photo by Yu. G. Men'shikov)

Figure 2. The carriage of a peacefully moving
monitor, (photo by A.Yu.Tsellarius)

1997

1W7

Asiatic Herpetological ResearchVol. 7, p. 125

Figure 3. Concealing himself, the monitor presses his body tightly to the ground (3a, above left) and remains
motionless. The animal's eyes remained open in every instance of this that we were able to make out. Convincedthat he has not been noticed, the monitor moves away from the site of the encounter after a short time,

creepingalong the ground such that his elbows and knees frequently rise higher than the level of his back (3b, above right).
The animal moving away from the observer in this fashion uses shrubs and elements of relief so skillfully as cover

Figure4.

The carriage of the monitor in flight. The animal is fleeing the observer, (photo by Yu. G. Men'shikov)

Figure 5. The male drags his body along the ground, flattening the substratum in his
wake (signaling marks, below transliterated from the Russian as "zatirs") in the area

which a female from his settlement (The Fourth) and males from a neighboring settle-

ment (Rhombik and Mafiozi) had visited not long before. This took place on June 5,
1993 in a region where the borders of neighboring settlements touch and animals fromboth settlements visit, (photo by A.Yu.Tsellarius)

Asiatic Herpetological ResearchVol. 7. p. 126

Figure6.

1997

The "sitting dog" posture. When the moni-

tor is moving away from a site where he had encoun-/W

tered danger, the more frequent display of thereference reaction "What's going on?" is quite appar-

The lizard periodically -- sometimes every few
steps
sharply "falls" on his rear, raises his head andent.

the frontal section of his body on erect forelegs, andlooks around, (photo by A.Yu.Tsellarius)

Figure 7. The display of readiness for active selfdefense characteristic of the monitor lizard facing athreat from a human or another large animal (7a, atleft, top). If an opponent does not start an active operations the monitor, continuing the display of readinessfor defence, slowly retreats (7b. at left, bottom),(photoby A.Yu.Tsellarius)

Ul

-a*

Figure 8. Finding himself in an unavoidable position, the monitor opens his mouth widely in the direction of thedanger (8a. above left). An attack very frequently follows this and the monitor seizes his enemy in a "mortal grip"(8b, above right), (photo by A.Yu.Tsellarius)

Vol. 7, p. 127

Asiatic Herpetological Research

1997

Figure9.

The male lifts a head moving away from thefemale which assumes the posture of submission (fordetail see description of encounter in section "Exam/-.'>'.""imi!&m\'ples of behavior...", ex 1). (photo by A.Yu.Tsellarius)"

Figure 10. Diagram of trails of encounter between male and female (for detail see ex 2).
4 - shrub.

1-

male's trail and placeof lying; 2 - zatir of mail; 3 - female's trail and place of lying,

Figure
ius)

1 1

.

Female "shows the back" to familiar male which goes out of burrow (see ex 3). (photo by A.Yu.Tsellar-

Asiatic Herpetological ResearchVol. 7. p. 12*

1997

trail of young male (aFigure 12. Diagram of trails of encounter between two males (see ex 4). 1
the run); 2 - trail of old male (a - place of basking, b - the run. c - "threatening gait").
-- fl «, «*, >y» U

4 %. *3fe*

r

F_

U"4

m *

ft
Figure 13a

Figure 13c

Figure 13b

Figure 13d-

the step, b

1997

Asiatic Herpetological ResearchVol. 7. p. 129

V* _ ».»
Figure 13e-

Z^JU^mM

Figure 13f

Figure 13 (a-g). Ritual combat between males (fordetail see ex 5). a - mutual "sniffing'; b, c - wrestling bynecks; d - "waltz" with "showing of the backs"; e - transition to standing position on the hind legs; f • wrestling in vertical position; g-loss of equilibrium, (photoby A.Yu.Tsellarius)

Figure 13g

Figure 14a

Figure 14b

The encounter of a settled male with athe nomad heads for the settler
by "confident gait"; b the nomad assumed a posture
Figure14.

nomad (see ex 7). a--of appeasement; c - the nomad "sniffs" the track of thesettler,

Figure 14c(photo by A.Yu.Tsellarius)

Vol. 7. p. 130

Figure 15a

1997

Asiatic Herpetological Research

Figure 15b

toFigure 15. The encounter of a settled male with a new settler (see ex 8). a stooping new settler approaches
the "host": b - new settler (above right) attempts to "sniff" the sacrum of "host", (photo by A.Yu.Tsellarius)-