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An ethogram for rhesus monkeys I. Antithetical contrasts in posture and movement

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An Ethogram for Rhesus Monkeys
I. ANTITHETICAL CONTRASTS IN POSTURE
AND MOVEMENT
DONALD STONE SADE
Caribbean Primate Research Center, University of Puerto Rico, P u n t a
Santiago, Puerto Rico 00742 a n d Department of Anthropology,
Northwestern University, Evanston, Illinois 60201
ABSTRACT
Antithetical postures and movements in social displays of rhesus
monkeys distinguish five moods. Movement from a high to a low position in the
mid-sagittal plane indicates threat. Emphasis on movement in the frontal plane
indicates subordinance. Movement from a low to a high position in the midsagittal plane indicates non-hostile, non-fearful approach. Oblique movements
indicate a querying mood. Playfulness is indicated by emphasis on rotatory
movements in the transverse plane.
Several investigators have produced ex- tration of the posture of a hostile dog prehaustive or partial catalogues of the com- paring to attack contrasted with that of
municative or expressive behavior of a fawning friendly dog exhibiting what
Mucaca mulatta (Hinde and Rowell, '62; we now would call submissive or subordiAltmann, '62a; Reynolds, n.d., cited in Alt- nate behavior. A second set of illustrations
mann, '68; Sade, '67, '71). These show portrayed a cat preparing to fight conconsiderable consistency and agreement trasted with the attitude of the same aniamong themselves and also with the rele- mal rubbing affectionately against the leg
vant portion of Van Hooffs ('67) study of of a human. Behaviorists have since found
facial expressions among the Cercopithe- that the principle of antithesis applies to
cidae. Altmann's ('68) comparison of sev- a wide variety of animal taxa. Antithesis
eral of these catalogues indicates that i n of displays reduces confusion between
spite of good general agreement there is communicative signals, particularly bestill considerable further analysis to be tween signals implying threat on the one
done before the relations between the vari- hand, and subordinance or appeasement
ous behavioral elements become clear.
on the other. Most users of the principle
This report does not attempt to list be- of antithesis have contrasted pairs of
havioral elements but rather describes communicative acts.
some contrasts between postures and
A consideration of the expressive posmovements which seem to characterize tures and movements of the rhesus monsome diverse moods of the rhesus monkey. key, insofar as they contrast in the direcThe report is based on observations of free- tion and orientation of movement within
ranging rhesus monkeys on Cay0 Santiago, the anatomical planes, indicates that a t
Puerto Rico, supplemented by examination least five moods may be distinguished by
of 8 mm film records taken at the same the antithetical characteristics of their
colony by the author and by James Loy.
outward expression.
THE PRINCIPLES OF DARWINIAN
ANTITHESIS
Darwin (1872) showed that moods
which contrast markedly are accompanied
by expressive behaviors which emphasize
opposite or strongly contrasting characteristics. His example consisted of a n illusAM. J. PHYS.ANTHROP., 38:
537-542.
T H E ANATOMICAL PLANES
The mid-sagittal plane divides a monkey
along its axis of bilateral symmetry. The
frontal or coronal plane lies perpendicular
to the mid-sagittal plane and parallel to
the longitudinal axis of the body or the
vertebral column. The transverse plane is
537
538
DONALD STONE SADE
perpendicular to both the frontal and sagittal planes and transects the longitudinal
axis of the body.
The mid-sagittal plane undoubtedly represents a very real and important axis of
self-reference for the animal in his orientations to his physical, biotic, and social
surroundings (Von Uexkull, '21). The frontal and transverse planes are convenient,
geometrically defined, reference loci for
the anatomist in describing the relations
of anatomical parts or positions, but they
do not correspond to anatomically defined
natural divisions of the body. Figure 1G
shows the relations of the anatomical
planes superimposed upon a rhesus monkey in a neutral attitude.
the activation of a particular mood are
minimal or absent, the animal's posture
and movement may be considered neutral
in respect to expressive behavior and the
animal may be said to be in a neutral
mood.
The descriptions which follow apply to
cases in which the masking effects of the
primary locomotor requirements of balance
and orientation are minimally present.
The sketches do not contain information
about the temporal patterning of the shifts
in posture. The sketches only indicate the
kinds of contrasts which are more easily
observed in life or on film, where the temporal patterning of the changes in pose
are also observable.
CONCEPT OF THE NEUTRAL MOOD
GENERAL AROUSAL
The neuro-musculo-skeletal system carries out the primary functions of balance
and locomotion. Evolution has made use
of this primary system by superimposing
upon it a repertoire of movements and
poses which serve the function of indicating the mood of the animal and communicating it in the visual channel to conspecifics. Usually the functions of balance
and locomotion will be manifest and take
precedence over the expressive functions.
The independence of the expressive functions in the organization of the animal's
behavior is indicated by occasions when a
highly aroused individual performs a communicative display with such intensity on
an inappropriate substrate, such as a narrow tree limb, that he loses his balance
and falls to the ground.
The neuro-musculo-skeletal system also
provides the mechanisms by which a n individual orients towards events, objects, or
other individuals in the environment. Orientation may involve a modification in
posture whether or not a mood which
normally is accompanied by expressive
behavior is also active at the moment.
Expressive movements may be altered
by the requirements of simultaneously activated locomotor or orienting movements.
Unusual or bizarre postures may result
when the display is directed toward a partially concealed individual, or toward one
who is vertically removed from the displaying monkey. Conversely, if the modifications of posture and locomotion due to
The transformation of a relaxed individual into a n aroused and alert one has
often been observed. The alert state has
correlates in facial expression which have
been described by Van Hooff ('67), and
in postdres which have been described by
Hinde and Rowel1 ('62). General arousal
or alertness can be distinguished from the
activation of the particular moods to be
discussed below.
MOVEMENTS OF ATTACK
An attack by a confident animal may
begin with a bobbing of the head accompanied by a n open mouth directed towards
the victim, grade into a lunging of the
shoulders toward the victim, and finally
become a charge ending when the dominant individual bites the victim. These
are some of the best known of the expressive movements of the rhesus monkey and
are recognized by all the above cited authors who have studied this species. Careful analysis of the behavior of the attacker
show that movements from a high to a
low position in the mid-sagittal plane are
emphasized (fig. lA,B). If a mildly
aroused attacker is not facing his opponent at the beginning of a fight, he may
begin by merely turning and bobbing his
head toward the victim. In the same
episode, however, if the victim does not respond satisfactorily, the attacker will usually adjust his posture so that the longitudinal axis of his body is oriented directly
towards the victim and the bobbing
I
E
Fig. 1 Poses of rhesus monkeys. Sequence A, B shows movement from a high to a low position i n the mid-sagittal plane
indicating threat. Pose C shows the lateral flexion in the frontal plane characteristic of a subordinate animal. Pose D shows
another animal displaying subordinance. Sequence E, F shows an animal giving an upward jerk in the mid-sagittal plane. Pose
G is a monkey in a neutral sitting posture upon whom the anatomical planes have been superimposed. Pose H shows one member of a wrestling pair rotating head and torso in the transverse plane. Sequence I, J, K, L shows the oblique bobbing movements
of a querying animal.
C
540
DONALD STONE SADE
movements now involve head, shoulders,
and thorax. The point here is that as the
attacker becomes increasingly aroused the
high to low movements in the mid-sagittal
plane become exaggerated.
MOVEMENTS OF SUBMISSION
The postures and movements of the
victim contrast strongly to those of the
attacker in that they emphasize lateral
flexion of the vertebral column and other
movements within the frontal plane. As
is well known, the victim may attempt to
present both his hind quarters and his
grimacing face towards the attacker simultaneously, so that the animal’s body is
displayed laterally to the attacker. If the
victim flees, the tendency for lateral flexion disappears, undoubtedly overridden by
the requirements of rapid and directed
locomotion. Nevertheless, a slight lateral
flexion in the frontal plane is almost always given by a subordinate threatened
by a dominant, if only for a fleeting second, and has given rise to such appellations as cringing or cowering (fig. lC,D).
AMICABLE, NON-HOSTILE, NON-FEARFUL
APPROACH BEHAVIOR
Figure lE,F shows a display which contrasts strongly with both the movements of
the threatening or attacking animal and
also with the movements of a frightened
subordinate. It consists of a monkey in a
neutral pose rapidly extending (dorsiflexing) its head and sometimes torso upwards
in the mid-sagittal plane. This movement
may be repeated in a series of jerks in
which the upward movement is strongly
emphasized in contrast to the return downward movement. At the extreme high position the head cannot be further extended
(or dorsiflexed). At this point, as if the
mood were frustrated by the limitations of
the cervical articulations, some rotatory
movements (not illustrated) take place as
the upward pointed chin strives to remove
itself even further from its horizontally directed position in the neutral posture. This
upward jerking display may frequently be
accompanied by protrusion and smacking
of the lips and a lateral and whipping
motion of the tail (also noted by Altmann,
’62a; Sade, ’71; Lindberg, ’71). This display in its mildest form may consist merely
of a slight elevating of the muzzle in the
mid-sagittal plane accompanied by protrusion of the lips. It is given on occasion by
adult males and females and juveniles of
each sex when approaching either a subordinate or a dominant individual. It is
often seen displayed by a n adult female
approaching another female with a young
infant. It is often directed towards a n infant, other than the female’s own, whom
the female attempts to retrieve or kidnap.
Since the display is so obviously antithetical to both those of a threatening and of
a frightened and subordinate animal, the
existence of an additional mood is suggested which has not been clearly pointed
out in the primatological literature. The
term “reassurance,” which I first used to
characterize this display, indicated my
interpretation that it conveyed a friendly
and non-agonistic attitude on the part of
the approaching animal. However, “reassurance” has previously been used by Ewer
(’68) to designate behavior which tends
to increase the self-confidence of the performer while simultaneously intimidating
a stranger: therefore the circumlocution
of the subheading.
QUERYING MOVEMENTS
Count (’69) pointed out that all communicative displays of vetrebrates are attempts to query the social environment
since they function to elicit a response
from a conspecific. Whereas some displays, such as threatening or submissive
gestures, seem to function to elicit specific
responses from another individual, other
displays seem to function to test the mood
of the second individual without necessarily bringing about any specific response.
The sequence shown by figure II,J,K,L
illustrates the bobbing movements often
directed by monkeys towards other monkeys, humans, other species, or novel
objects. The open mouth face of the monkey illustrated in figure 11,K indicates the
presence of an aggressive component in
this particular case. However, other examples would show that the aggressive component may be diminished, lacking, or
replaced by some other mood. The aspect
of this display which is antithetical to the
previously discussed displays is the rapid
alternation of downward and upward
A N ETHOGRAM FOR RHESUS MONKEYS
movements and especially the emphasis
on oblique movements of the head, which
crosses the mid-sagittal plane as it is lowered and raised. An animal who alternates
rapidly between attack and flight shows
movements which have little in common
with the oblique bobbing display. An animal so motivated switches rapidly from the
mid-sagittal attacking movements to the
laterally flexed cowering movements of
submission, A subordinate animal attacking a dominant likewise displays postures
and facial expressions which are clearly
resultants of the simultaneous activation
of attack and flight tendencies (defensive
threat) and which bear little similarity to
the oblique movements discussed here. The
antithesis of these oblique and erratic bobbing movements to the other displays discussed above suggests that a different
mood is active, which I call the “querying
mood.”
PLAY
Any normal movement may be exaggerated during episodes of play among monkeys and play behavior is often interpreted
a s representing a n incomplete or distorted
form of behavior characteristic of some
other mood (Meyer-Holzapfel, ’56). However, observation of play episodes among
rhesus monkeys indicates that a postural
component is often present which seems
to be unique to play behavior, namely rotation of the head or torso in the transverse
plane (fig. 1H). An animal running towards a play episode or i n the chases
which commonly occur between bouts of
wrestling usually is recognized by its peculiarly bouncing or gamboling locomotion.
Careful observation indicates that this type
of locomotion differs from a neutral, aggressive, or subordinate type in part by
the tendency of the playfully running animal to rotate its head and even shoulders
and to abduct its limbs during the noncontact phases of the gait. This produces
the impression of movement around the
longitudinal axis of the body, or movement
in the transverse plane. It may be that
this tendency for movement in the transverse plane is the “metacommunicative”
message which Altmann (’62b) postulated
a s being necessary for play to be distinguished from activities with more serious
54 1
consequence. Another interpretation could
be that because the movements of play are
clearly antithetical to those of the other
moods discussed above, play should be
considered a mood in its own right and
not simply derived from other aspects of
behavior.
COMPARATIVE
Limitations of space preclude detailed
comparison between this study and other
reports. However, examination of photographs of Papio hamadryas (Kummer, ’68).
Theropithecus gelada (Spivak, ’68). Miopithecus talapoin (Gautier-Hion, ’71 ) and
unpublished personal observation on Cercopithecus aethiops indicate that these
postural characteristics may be widespread
in the Cercopithecidae.
ACKNOWLEDGMENTS
This work was supported by NSF grant
GS-3114, and by Contract NIH 71-2003
from NINDS, NIH, HEW. Conversations
with Glenn Hausfater in 1968 clarified
some aspects of the displays described in
this paper. Comments by Judith Breuggeman, Jane Buikstra, B. Diane Chepko-Sade,
and Carol DeRousseau helped clarify obscurities in the manuscript. Mr. Hemlock
made the sketches. Any faults or errors
are the author’s.
LITERATURE CITED
Altmann, S. A. 1962a A field study of the
sociobiology of rhesus monkeys, Mncaca muZatta. Ann. N.Y. Acad. Sci., 102: 338-435.
1962b Social behavior of anthropoid
primates: Analysis of recent concepts. In:
Roots of Behavior. Chap. 20. E. L. Bliss, ed.
Harper, New York, New York, pp. 277-285.
1965 Sociobiology of rhesus monkeys.
11. Stochastics of social communication. J.
Theoret. Biol., 8: 490-522.
1968 Primates. I n : Animal Communication. Chap. 18. T. A. Sebeok, ed. Indiana
University Press, Bloomington, pp. 466-522.
Count, E. W. 1969 Animal communication i n
man-science: An essay i n perspective. In: Approaches to Animal Communication. Chap. 4.
T. A. Sebeok and A. Ramsay, eds. Mouton &
Co., The Hague, pp. 71-130.
Darwin, C. 1872 The Expression of the Emotions in Man and Animals. University of Chicago Press, Chicago, pp. 1-366.
Ewer, R. F. 1968 Ethology of Mammals. Logos
Press Limited, Great Britain.
Gautier-Hion, A. 1971 Repertoire comportemental du Talapoin (Miopithecus talapoin). Ext.
Rev. Biol. Gabon., 7: 295-391.
542
DONALD STONE SADE
Hinde, R. A., and T. E. Rowel1 1962 Communication by postures and facial expressions in
the rhesus monkey (Macaca mulatta). Proc.
Zool. SOC.Lond., 138: 1-21.
Xummer, H. 1968 Social Organization of Hamadryas Baboons. A field study. University of
Chicago Press, Chicago.
Lindberg, D. G. 1971 The rhesus monkey in
North India. An ecological and behavioral
study. In: Primate Behavior. Vol. 11. L. A.
Rosenblum, ed. Academic Press, New York,
pp. 1-106.
Meyer-Holzapfel. M. 1956 Das Spiel bei Saugetieren. Handb. d. Zool., 8: 1-36.
Reynolds, V. n.d. The social behaviour repertoire
of rhesus monkeys (Macaca mulatta) in a
captive colony.
Sade, D. S. 1967 Determinants of dominance
in a group of free-ranging rhesus monkeys. In:
Social Communication Among Primates. Chap,
7. S. A. Altmann, ed. University of Chicago
Press, Chicago, pp. 99-114.
1971 Communication by tail positions
in rhesus monkeys (Macaca mulatta). Am. J.
Phys. Anthrop., 35: 294. (Abstract)
Spivak, H. 1968 Ausdrucksformen und soziale
Beziehungen in einer Dschelada-Gruppe (Theropithecus gelada) im Zoo. Juris Druck & Verlag, Ziirich.
Van Hooff, J. A. R. A. M. 1967 The facial displays of the Catarrhine monkeys and apes. In:
Primate Ethology. Chap. 11. D. Morris, ed. Aldine Publishing Co., Chicago, pp. 7-68.
Von Uexkull, J. 1921 Umwelt und Innenwelt
der Tiere (2nd Edition). Springer-Verlag, Berlin.
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