Primate Info Net Banner Wisconsin PRC Logo

Gorilla Ethograms

                    Collection of Gorilla Ethograms

                             Compiled by:

                    The Gorilla Behavior Advisory Group
                     [affiliated with the Gorilla SSP]

      Jackie Ogden, Zoo Atlanta and Georgia Institute of Technology
           Deborah Schildkraut, Ph.D., Boston MetroParks Zoos
                                Co-chairs

                    Anne Baker, Ph.D., Brookfield Zoo
                  Benjamin B. Beck, Ph.D., National Zoo
                   Cynthia Bennett, Ph.D., Dallas Zoo
                      Thaya duBois, Los Angeles Zoo
                  Cathleen Cox, Ph.D., Los Angeles Zoo
        Ken Gold, Zoo Atlanta and Georgia Institute of Technology
             Donna Fernandes, Ph.D., MetroParks Zoos, Boston
                  Carol Glick, Arizona State University
                      Ingrid Porton, St. Louis Zoo
           Jill Mellen, Ph.D., Washington Park Zoo, Consultant



The Gorilla Behavior Advisory Group (GBAG) would like to gratefully
acknowledge all those researchers who gave permission to include their
ethograms in this compilation, as well as those who had previously
published their ethograms.  Additionally, GBAG would like to thank the
Gorilla SSP Group for their sanction.  Each ethogram is listed under the
name of the respective author(s), with either the author's affiliation or
the reference from which it was taken.  The majority of these ethograms
were, of course, built upon previous work of other researchers.  Any
typographical errors or errors of interpretation are of course the
responsibility of the compilers, not the respective authors.  This work
represents a compilation, not a recommendation of any particular
ethogram.  Although an attempt was made, this compilation is by no means
exhaustive.  Copies of this compilation are available for $5.00 from
Jackie Ogden, Zoo Atlanta, 800 Cherokee Avenue SE, Atlanta, GA  30315. 
[Checks should be made payable to the Atlanta/Fulton County Zoo.]  You may
also obtain a copy of this document in electronic form in either
WordPerfect 4.2 or in ASCII text-files [please indicate which version you
can use].  Send one formatted 5 1/4 inch diskette to Donna Hardy, Dept. of
Psychology, Calif. State Univ. Northridge, Northridge, CA 91330.  [Since
this is a service of the Consortium of Aquariums, Universities and Zoos,
no charge is made for the ethograms in electronic form.]

                          updated January, 1991







                            TABLE OF CONTENTS

Beck, B. Phase 1...............................................page 3
Beck, B. Phase 2...............................................page 5
Bennett, C. and Fried, J. (1990)...............................page 7
Bowen, R.A. (1980).............................................page 25
Brown, S.G. (1988).............................................page 26
duBois, T., Mead, J. & Cox, C..................................page 27
Elliott, R.C. (1976)...........................................page 37
Fischer, R.B. (1984)...........................................page 38
Fossey, D. (1972)..............................................page 39
Fossey, D. (1979)..............................................page 41
Fossey, D. (1982)..............................................page 45
Frisch, D. J...................................................page 46
Goerke, B., Fleming, L., & Creel, M. (1987)....................page 48
Gold, K. (a)...................................................page 50
Gold, K. (b)...................................................page 52
Gould, J. E....................................................page 54
Harcourt, A. H. (1979a)........................................page 67
Harcourt, A. H. (1979b)........................................page 68
Harcourt, A. H. & Stewart, K. J. (1981)........................page 69
Hoff, M. P., Nadler, R. D., & Maple, T. L. (1981a).............page 70
Hoff, M. P., Nadler, R. D., & Maple, T. L. (1981b).............page 72
Lockard, J. S..................................................page 73
Mannchen, K.  (1980)...........................................page 79
Maple, T.L. & Hoff, M..........................................page 81
Meder, A. (1985)...............................................page 86
Meder, A. (1986)...............................................page 87
Meder, A. (1990)...............................................page 88
Mitchell, R.W. (1989)..........................................page 89
Nadler, R.D., Collins, D.C., Miller, C., & Graham, C.E. (1983).page 90
Nadler, R.D. (1985)............................................page 91
Ogden, J.J., Hoff, M., & Maple, T.L............................page 92
Quiatt, D., Miller, L., & Cambre, R. (1986)....................page 96
Riess, B.F., Ross, S., Lyerly, S.B., & Birch, H.G. (1949)......page 97
Schaller, G. (1963)............................................page 99
Schildkraut, D. & Akers, J.....................................page 100
Schildkraut, D. (a)............................................page 102
Schildkraut, D. (b)............................................page 103
Schildkraut, D. (c)............................................page 105
Schildkraut, D. (d)............................................page 106
Schildkraut, D. (e)............................................page 107
Smalley, S.L. (1979)...........................................page 108
Watts, D.P. (1988).............................................page 109
Watts, D.P. (1990).............................................page 110
Woods, S.......................................................page 111
Wood, G., Forthman, D.L., & Ogden, J.J.........................page 113
Yamagiwa, J. (1986)............................................page 115
Index..........................................................page 116
Appendix.......................................................page 117




Beck, B.  Phase 1: National Zoo, Gorilla Research Project 
          (study at the Brookfield Zoo)

Purpose: To compile data on the social behavior and social structure of
gorilla groupings to better inform management decisions and to plan
strategies for reproduction.

Sampling methods: Scans of location and activity (5 minute intervals).  
Other behaviors, all occurrences.

Ethogram key words: Activity budgets/General behaviors, Social behavior.

Behaviors:

Activity:

Lie: Being recumbent with little or no weight being borne on hands or
feet.

Sit: Bearing most weight on buttocks.

Stand still: Bearing weight on feet quadrupedally or bipedally with
hands holding on to mesh, glass, tree, etc.

Locomote: Walking, running, or climbing.

?: If activity can't be scored.

Other behaviors:

Rest Near: Sitting or lying within arm's reach of another for at
least 10 seconds.  Initiator is second animal to rest.  The receiver
must be relatively immobile but need not be resting.

Groom: Directed touching, licking, or intense visual inspection of
another's (GO) or own skin (GS) or pelage.  Scratching excluded.

Intimidation display: Resting on chest or abdomen, beating on cage
surface, rapid exaggerated running or strutting, pursed lips, and/or
exaggerated throwing or shoving of objects.  Score when any two or
more of these behaviors occur simultaneously or sequentially.  Score
initiator only.

Rough-up: Slapping, poking, elbowing, punching, hair pulling,
pushing, shoving, or wrestling when not in play or sexual context.

Aggressive bite: Biting when not in play.

Crouch: Initiator lowers ventrum toward or to substrate, or curls
fetally on substrate.  Event terminated when normal locomotor or
resting posture resumed.

Inspect genitalia: Sniffing, touching, or intense visual inspection
of another's genitalia or own genitalia.

Sex solicit: Female purses lips and stares at/follows/reaches toward
male.  Event terminated when lips relax for at least 10 seconds.

Copulation: Dorso-ventral mounting or ventro-ventral mounting with
pelvic thrusting.  Event terminated when either animal breaks bodily
contact.  Note if intromission confirmed (very difficult) or if semen
observed afterward on penis or vagina.

Masturbate: Rhythmic rubbing of genitalia with any part of own body
(e.g., fingers, toe), or against inanimate object.

Yawn: Per usual definition.  Score each yawn.

Feces: Manipulation or mouthing of own or another's feces.  Note
whose feces if possible.  Note "eat" if fecal material actually
ingested.

Regurgitate: Vomiting into mouth, hand, or onto substrate; note if
reswallowed.

Termination of behavior: Termination of yawn, crouch, sex solicit and
copulation is specified in definition of those patterns.  For all
other patterns, an event is terminated when discontinued for 10 or
more seconds or if the participating animal(s) engage in another
scored pattern within 10 seconds.  However, Rest Near is not rescored
until one partner moves out of arm's reach for 10 seconds regardless
of concomitant behavior. 


Beck, B.  Phase 2: National Zoo, Gorilla Research Project 
          (study at the Brookfield Zoo)

Purpose: To study social development of captive-born lowland gorilla
infants being mother-raised in a group.

Sampling methods: 1-0 sampling at 60 seconds intervals.  Note occurrence
of rough-up, aggressive bite, or crouch by putting initiator-receiver
letter codes. 

Ethogram key words: Development.

Behaviors:

Infant location relative to other animals:

Contact: In physical contact with mother, whether carried or simply
touching.

Other contact: Any physical contact between infant and nonmother.

Proximate: Within arm's reach of other animal, but only when not in
contact with mother.

Distant: Beyond arm's reach but less than 15 feet of other animal,
but only when not in contact with mother.

Independence:

Nursing: Infant's mouth on mother's nipple, with or without actual
suckling.

Eating: Infant puts any solid food (including hay) into its own
mouth, with or without actual chewing or swallowing.

Wean: Mother actively terminates or prevents nursing.

Struggle: Infant trying to break contact withmother while being
restrained by her.

Play :Low intensity slapping, wrestling, or mouthing when initiator
has slack lips or "smile."

Aggressive behavior:

Rough-up: Slapping, poking, elbowing, punching, hair pulling,
holding, or wrestling, when not in play.

Aggressive bite: Biting, when not in play.

Crouch: Initiator lowers ventrum toward or to substrate, or curls
fetally on substrate.

Copulation: Dorso-ventral mounting or ventro-ventral mounting with
pelvic thrusting.  Event terminated when either animal breaks bodily
contact.  Note if intromission confirmed (very difficult) or if semen
observed afterward on penis or vagina.

Masturbate: Rhythmic rubbing of genitalia with any part of own body
(e.g., fingers, toe), or against inanimate object.


Bennett, C. and Fried, J. (1990)  Dallas Zoo.  Gorilla Ethology Study.

Purpose: To investigate the effects of exhibit modifications on gorilla
activity budgets.

Sampling: 15 minute focal sampling, with scan sampling at 15 minute
intervals of all individuals.

Ethogram key words: Activity budgets/General behaviors, Social behavior,
Sexual behavior, Post-occupancy evaluation, Display/Agonistic behavior,
Abnormal/Stereotypic behavior.

Behaviors:         

Locomotion:

Walk: Individual travels from one location to another by using two,
three, or four limbs to propel the body.  Two or three limbs are
always on the ground at one time. 

Walk quadrupedal :The animal uses all four limbs to travel forwards,
backwards, or to the side.  Leg movement is transverse (i.e., left
forefoot and right hindfoot moving together, then right forefoot and
left hindfoot moving together).  Gorillas primarily knucklewalk, but
occasionally walk on the sides of closed fists.  

Walk bidpedal: In this form of locomotion, the torso is perpendicular
to the ground.  Travel is executed by using hind limbs alone.  Legs
are extended and support the body.  Forward motion of the legs comes
more from a forward and backward rocking of the pelvis rather than
rotational movements in the hip joints.  Arms are usually held close
to the side or are wrapped around the torso.  Sometimes, individuals
walk bipedally for short distances while holding onto a structure
(i.e., cage bars, bench, door frame edge, ropes, trees, vines) or
while carrying object(s).

Run: Three gaits can be observed: transverse quadrupedal, lateral
quadrupedal (gallop) and bipedal.  Regardless of the gait; none, only
one, or two limbs will be on the ground at a time.  Bipedal run is
achieved by slightly elevating the torso and lifting the arms during
a transverse run.  This usually occurs within aggressive display
sequences or bouts of play behavior.

Slide: Occurs at end of a run.  The animal remains quadrupedal but
stops the running motion while keeping the limbs rigid.  The body
might be turned at an angle to the direction of travel.  A slide may
be terminated by the presence of cage bars, wall, limbs, etc.

Arm over arm swing: An animal grasps an overhead structure with its
hands.  The arms are nearly fully or fully extended.  The animal
releases one hand and pivots the body forward from the still grasping
hand while extending the other hand to grasp a structure.  It then
pivots the body forward.  Swinging movements are slow and deliberate. 
Reminiscent of brachiation.

Skip: Not observed in Dallas animals.

Swing: Not observed in Dallas animals.

Circle: Not observed in Dallas animals.

Scoot: Two forms of scoot have been observed, and both occur with the
animal in the squat position.  One is a wobble like movement, where
the animal leans from side to side while swinging the elevated side
of the body forward.  This can be done with the arms crossed on the
chest or resting on the knees, but the arms could be used to pull the
animal along while it moves next to the cage bars or ropes.  The
second form is characterized by use of the arms in a crutch like
manner.  The arms are placed on the ground in front of the body. 
With the arms rigid and braced on the ground by tightly closed
fists, the body is pivoted forward by swinging between the arms. 
Both modes of scoot are primarily used for short distance travel, but
individuals have been observed to travel up to twelve feet in this
manner. 

Climb: Gorillas are methodical climbers moving carefully and always
having two limbs in contact with a structure at any one time.  The
type of structure being climbed appears to influence the specific
climbing movements used.  When climbing cage bars, wood ties, or
trees, movement of the limbs is basically transverse quadrupedal
(transverse limbs may move together or in series).  When climbing up
ropes or horizontal bars, an animal may grasp them with its forelimbs
and partially swing its body up off of the ground until the hindlimbs
can also grasp or lean on the structure, thereby assisting the
forelimbs in pulling the body up.

Inactive:           

Sit: Basically, sitting consists of: torso erect, not in contact
with ground, resting platform, substrate, etc.; buttock on ground,
legs in front or to the side of the body.  Several variations of
sitting postures are regularly displayed: 
     Legs: bent acutely at the knees, rotated outwards from body
(outer thighs are close to the ground), and crossed at the ankles or
shins (aka: Indian style).  The legs can also be extended forward and
crossed at ankles, bent with knee(s) up and foot/feet flat on ground,
legs extended and near parallel, or extended and split apart up to a
90 degree angle.
     Arms/hands: crossed on chest, crossed or lying in lap, gripping
feet, resting on knee(s), or extended with knuckles resting on ground
for additional support.  Animals will frequently hold on to structure
(i.e., ropes, cage bars, cross-ties) or rest limbs on it. 
     Torso: upright almost perpendicular to the floor, angled acutely
forward over legs at about a 45 degree angle or less to the floor, or
leaning to the side supported by structure, object, or extended arm. 
Record as torso upright, torso hunched over, or back, side or arms
supported.

Squat: In this posture, legs are acutely bent at the knees and
thighs, with the feet under the body and heels tucked into the
buttocks.  Feet and legs bear the bulk of the weight and can be in
front of the body or to the side.  Buttock does not contact surface. 
A squat position is assumed when the animals defecate or urinate (see
Elimination category).  There are several variations of the squat
position:
     Legs -  Knees are bent and can be tucked up in front of the
chest or angled outwards until they form an almost straight line with
the body.
     Arms - Forearms or elbows are frequently resting on knees,
thighs or holding onto a structure (i.e., cagebars), but may also be
crossed on chest or extended to the ground for additional support.
     Torso - Squatting animals will frequently have their backs
upright against a structure (i.e., cage bars, walls, concrete posts). 
When supported by arms extended out, the torso may be bent at a 10
to 15 degree angle.  

Perch: Animals assume a squat posture when they are off of the
ground.  The feet are used for grasping surfaces.  
     Legs/feet - Grasp structure with feet side by side or one in
front of the other. 
     Arms/hands - Grasp same structure/object or nearby
structure/object with hand(s) for additional support.

Lie: In this posture, the body is in a recumbent, prostrate
position.  The body can be dorsally, ventrally, or laterally
recumbent.  Variations within these positions include:
     Ventral: Primarily the ventrum is on the resting surface. 
     Variations of ventrum include:     
          Arm(s) - May rest chin on forearm or hand.
          Legs - May be bent at knees and tucked the body or hips may
          be rotated so both legs lay out to the side.
     Dorsal: Primarily the back is on the resting surface. 
     Variations of dorsal include:
          Arms/hands - Are frequently folded across the chest or are
          elevated with the elbows extended out and hands resting on
          the head.  Occasionally, the arms are extended outwards and
          are grasping a structure.
          Legs/feet - May be crossed at ankles or knees, sometimes an
          ankle may rest on opposite knee.  The feet may be gripping
          an object or could be propped up against a structure. 
          Sometimes, legs may just stick up in the air at a  90
          degree angle to the plane of body.  Foot-foot and
          hand-foot grasping occurs frequently.
     Lateral:  Primarily the left or right side of the torso is on
     resting surface.  Variations include:
          Arms/hands - May be folded on chest, lying along length of
          body or one/both arm(s) may be bent up under head as a
          "pillow."
          Legs/feet - Are usually bent at knees, pulled up towards
          the body, and laid one on top of the other.  The legs also
          can be crossed at knees or ankles.  Foot-foot and hand-foot
          grasping occurs frequently. 
          Torso - The back may be against an object or unsupported.

Stand: Outstretched or extended arms and/or legs are in contact with
a surface and bear body weight.  Stand, like sit and lie, can take
several forms.
     Stand quadrupedally: All four limbs are extended, are in contact
     with a surface, and support the body weight.  The torso is
     generally parallel to the ground.  Animals will stand on
     knuckles or on closed fists; on flat feet with thumbs extended
     or on the side of the feet.  Variations on the basic quadrupedal
     stance can be identified as threat behavior (see descriptions on
     agonistic behavior).
     Stand bipedally: The torso is in an upright or nearly upright
     position (perpendicular to the ground).  The legs support the
     body, are widely spaced, and bowed or slightly bent.  The feet
     are turned out, and thumbs extended out from foot.  There is no
     other support.
     Stand supported: The animal is in a bipedal or monopedal
     position with the torso perpendicular or even parallel to the
     ground.  The weight is supported by the legs, hands, or arms,
     which may be grasping or resting on some structure. 

Environmental exploration:

Survey:           
     Visual: Animal stops an ongoing behavior or is engaged in no
     other behavior and orients head and eyes toward a specific
     direction or component of the environment.  This orientation can
     be prolonged (greater than 2 sec). 
     Visual side glance: The animal repeatedly shifts its eyes to the
     side and focuses momentarily on components of the environment. 
     The head is stationary and may be oriented away from the
     object(s) of scrutiny.
     Inspect: Includes both tactile and olfactory components.  The
     animal reaches out and touches an object with its knuckle(s) or
     forefingers.  The digit(s) are then brought to the nose and
     apparently sniffed.
     Olfactory: An animal brings the nose close to an object or the
     object near the nose and sniffs.
     Gustatory: An animal licks an object, places it momentarily in
     the mouth or between the lips, or touches something then places
     fingers in the mouth.

Object manipulation:
     Object-gather: An animal accumulates objects that are dispersed
     and brings them to a central location.
     Object-dissemble: Involves a wide variety of actions where the
     hands, feet, and mouth, are used to rip, shred, twist apart,
     remove components, or peel an object (normally inanimate, but
     not necessarily).
     Object hold: An animal is inactive/idle with an object resting
     in the hand, foot, mouth, or pressed against the body.  No other
     manipulation is occurring.
     Object carry: An animal uses its hands, mouth, legs, and/or feet
     to transport an object.  Objects may also be carried by tucking
     them between the thigh and belly.  
     Object rigid manipulation: An animal uses an object to rattle,
     shake, bang, or pound.
     Object use: Involves utilizing objects which are not permanent
     fixtures for a number of different purposes: use of an object as
     an extension of the body in order to attain an immediate goal
     (tool), to increase comfort (furniture, nest), as covering, as a
     weapon, etc.
     Object varied or non-specific manipulation: This is a catch all
     category that includes touching, pushing, absently rocking,
     bending, lip rolling, etc.

Auto involved: Manipulate, eliminate, play, ingest:

Manipulate:       
Autogroom: Animal uses fingers and/or lips and teeth to manipulate
and remove items from its fur.
Autopick: Animal uses fingers and/or lips to pick debris from its
teeth, nails, nose, ears, and eyes.
Probe anus: animal uses fingers to poke at or rub its anal region.
Masturbate: Self-stimulation of genitalia.
Manual: Animal manipulates its genitals with its hands or feet.
Pelvic rub: Female backs up to an object and rubs anogenital area
against it.
Haunch bounce: From a squatting position animal bounces up and down
on haunches.  Males may create a false vagina with hands and haunch
bounce while directing the penis into the hands.
Autoinspect: Animal makes a close visual or olfactory examination of
own body parts.
Comfort: Comfort movements consist of brief, transitory movements
such as changes in posture; e.g., rolling over and changing
orientation of limbs and attempts to ease or relieve an uncomfortable
situation; e.g., stretching, scratching, yawning, coughing, clearing
the throat, etc.
Eliminate:        
     Urinate: Expel urine.  Animals assume a squat position with
     torso tipped forward and weight on its arms which are extended
     in front of the body.
     Defecate: Expel feces.  Position during defecation is similar to
     that described for urinate.  Animals may urinate and defecate at
     the same time.
Regurgitate: To eject part or all of the stomach contents through the
mouth, usually in a series of involuntary spasms.  However, gorillas
seem to have a high degree of control over regurgitation.  Record
this only if the animal regurgitates but does not reingest vomitus.
Play: Lone play includes repetitive, exaggerated, disjointed, and
seemingly nonpurposeful behavior.  When alone, such behavior usually
involves an inanimate object or consists of repetitive movements
accompanied by a play face.
Object: Shake, bounce, hit, juggle item, hug, rub, swing, toss in
air.  Movement is exaggerated.  Water can be considered to be an
object.
Peregration or locomotor play: Clap, rock, somersault, shake, twist,
run, slap, pat, circle, lip flap.
Body oriented: Manipulate body part, exaggerated head and/or body
rubbing, pulling of extremities, etc. - chest beat.

Ingest: Includes a wide array of behavior associated with the
location, procurement, processing, and consumption of items.
     Forage: Manipulation of substrate or objects in the environment
     which results in obtaining food items (substrate is frequently
     edible itself).  This includes the periodic transfer of food
     from hand/foot to mouth and travel of one or two steps between
     searches.  Several foraging motions can be used.  For example,
     animals may sweep an arm from side to side and move substrate
     with the back of the hand or pick up the substrate and place or
     throw it aside, or a substrate might be picked up and held while
     an animal forages through it.  To pick up food items, animals
     will use a finger pinch (forefinger or middle finger and thumb
     or tip of forefinger or middle finger for small forage
     items).  A variety of postures may also be assumed while an
     animal forages: tripedal with free arm foraging, sitting,
     squatting, or lying ventrally while resting on forearm(s) or
     elbow(s).
     Food carry: Travel while holding food items.  Food can be
     carried by mouth, hand, feet, in the crook of the arm, against
     the chest, against the stomach, or any combination of these
     methods.  Animals walk bipedally (rarely), tripedally (most
     commonly), or quadrupedally.  Animals will knuckle walk with
     food in their hands or will walk on the back of the hands/wrists
     while holding food.  When climbing, animals will frequently
     carry food in their mouths.  Animals may also scoot along for
     short distances while carrying food.
     Food gather: See object gather.
     Food process: Various manipulations of food items (provisions,
     browse, bedding) that give an animal access to an otherwise
     inaccessible edible or preferred portion.  This might involve
     one or more of the following: stripping (usually length wise),
     peeling, shredding, breaking apart, husking, shelling, or
     shaking apart.  Each action can be carried out with the teeth,
     hands, lips, or feet, but usually occur in combination.  Visual
     and olfactory inspection of food frequently occurs during
     processing.  Individuals usually sit while food processing, but
     will also recline ventrally.
     Food beg: The animals will extend arms and clap hands together
     several times or will clap hands together and then open hands
     palms upwards.  Hands are held in a relaxed manner with fingers
     curled inwards.  Knuckles on each hand are actually tapped
     gently together.  If this does not get a response, the animals
     may rap on the bars or doors.  Grunt-like  vocalizations may
     accompany begging.  This behavior is usually  directed towards a
     keeper.
     Eat: Placement of food items in the mouth by hands, feet, or
     lips followed by chewing and swallowing.  "Eat" does not require
     that a substrate be manipulated in order to obtain food.  Visual
     inspection, olfactory inspection, manipulation, and/or food
     processing may occur prior to and during eating.  Food may be
     regular provisions (record separately), browse or forage (record
     separately), or body waste (record separately) (excrement,
     cerumen, nasal mucus, scabs, sweat, dead skin, etc). 
     Individuals have also been observed to wipe sweat from their arm
     pits with their hands and then lick their hands.
     Drink: Animals drink from fountains, pools, streams, or from
     areas where fluid has accumulated.  Similar drinking postures
     are used by the Dallas individuals.  From a quadruped stance, an
     animal either bends forward and places both hands against a
     wall, fountain edge, or on each side of the body; or crouches
     down with torso supported by all four bent limbs and ventrum
     close to the ground.  The head is horizontal when drinking.  The
     animals will lick up fluids or suck them directly into the
     mouth.  Fluids may be water or juice (record separately), or
     urine (record separately),
     Regurgitate and reingest: Expelling stomach contents through the
     mouth and then consuming the regurgitate.  Specify body position
     during regurgitate, location regurgitation is deposited in,
     method of reingest, body position during reingest.  Specify body
     position while regurgitating:
          Tilt position - The legs are straight or slightly bent at
          the knees.  Body is bent at the hips with the torso
          straight and angled downwards.  Body weight is supported by
          one forearm, both forearms (parallel along the orientation
          of the body, parallel and running across the chest or
          crossed across the chest) or one or both arms can be
          resting with the palms flat on the ground and elbows
          pointing out to the sides or backwards along the body. 
          The head is lower than the hips.  Dallas gorillas will
          regurgitate onto a surface or into their mouths from this
          position.  The gorillas will assume a tilt posture usually
          from a sitting or squatting stance.  Several head bobs
          frequently occur prior to regurgitation.
          Lie ventral - The animal reclines ventrally on the ground,
          resting on the forearms (which are held parallel across the
          chest).  Head bobs usually precede regurgitation.
          Quadrupedal stance - The animal assumes a quadrupedal
          stance with legs and arms bent to varying degrees, but the
          torso remains parallel to the ground.  Vomitus is
          forcefully expelled from 1/3 to 1/2 m (1.5 ft) above the
          receiving surface.
          Quick tilt - From a squat or sit position the animal
          rapidly tilts forwards on its palms then quickly returns to
          the previous position (looks like a pushup from a sitting
          position).  The gorilla might straighten its back legs a
          bit when tilting forward but bends them again when it
          returns to the sit/squat position.  Regurgitation is
          usually into the mouth and reingestion occurs while in the
          sit/squat position.
          Headshake - From a sitting or squatting position the
          gorilla moves its head to one side and slightly upwards in
          a somewhat circular motion or quickly up and down. The
          mouth might open and then close quickly.  The movement can
          be so slight that it may seem as if the gorilla is moving
          its head in order to relieve a crick in the neck.  It may
          be mistaken for a comfort movement.  The gorilla appears to
          regurgitate into the mouth; cheek bulging and chewing
          movement can be seen.  Vomitus is usually retained in the
          mouth.
     Methods of reingestion:
          Reingest with the hand - An animal will dip or run its
          fingers and whole hand into vomitus and then lick the
          material off.  Usually, the material has a liquid
          consistency.  An animal sometimes uses its hand to scoop
          and maintain vomitus in small puddle.   If the vomitus is  
          thick or chunky, the fingers and thumb, in a pincer motion,
          are used to pick up matter and transfer it to the mouth.
          Reingest with the mouth - Use mouth, lips and tongue to
          suck, lick, or pick up vomitus.  
     Positions for reingestion:
          Squat on elbows - Feet are flat on the ground, the body is
          bent forward at the hips so that the ventrum is on the
          ground.  The gorilla is resting on the forearm(s).  Hand or
          mouth will be used to reingest from this position.         
          Sit/squat - Animal assumes a sit or squat posture
          (described elsewhere) after the quadrupedal, quick tilt, or
          head shake regurgitation.
          Lie ventral - Similar to the squat on elbows position but
          the feet are not flat on the ground but extend out from the
          body.  The usually rests on the forearms. 
          Tilt position - This is the same posture as used during the
          regurgitation phase.  
Stereotypic, unusual, aberrant behavior: 
     Stereotypic: Stereotypic behavior is characterized by  excessive
     repetition of or lack of variation in vocalizations, movements,
     postures, or patterns of travel.
     Stereotypic posture: Prolonged maintenance of an unusual or
     bizarre stance, placement of extremities, or attitude.  Specify
     act.
     Stereotypic movement: Involves body or limb movements but not
     travel.
     Head circle: A series of 2-12 unidirectional head rotations
     which occur most frequently while in a quadrupedal stance with
     the head, neck, and shoulders oriented towards a corner of an
     enclosure.  The head is only 1/3 to 1/2 m from the wall.  This
     behavior has been observed only in the younger silverbacked
     male.
     Head swing: This motion is similar to head circles but the head
     hangs down and swings back and forth.  The head does not make
     full rotations.  It occurs mainly while sideways walking and
     occasionally while facing the wall or into a corner.  This
     behavior has been observed only in the younger silverbacked
     male.
     Rock: Legs are bent and the knees are near chest, arms are on
     knees or folded across chest.  From this position, the animal
     sways from side to side (movements are no more than 5-10 cm to
     each side).  This is usually done while the animal's back or
     side is against a vertical structure but has occurred while it
     is gripping the cage bars with its hands or while perched
     on a horizontal bar.  Rocking occurs in association with finger
     sucking.  This behavior has been observed only in the younger
     adult female.
     Hand clap: Hands, with fingers curled in, are brought together
     rapidly several times.  This behavior has been observed only in
     the younger adult female.
     Stereotypic locomotion: Repetitive, unvarying, and seemingly
     meaningless pattern of travel.
     Pace: Repetitious, patterned and usually unidirectional movement
     around the exhibit/habitat.  The animal seldom makes more than
     two complete circles about an enclosure.  Pacing speed is faster
     than the usual walking speed.  This behavior is seen in the
     younger silverbacked male and in both adult females.
     Stereotypic self-oriented: Excessive repetition of an unvarying
     form of self manipulation or self grooming.  
     Finger sucking: Animal will insert forefinger and/or middle
     finger of either hand completely into mouth.  Seen frequently in
     association with rocking.  Occurs while the animal is in a
     sitting, squatting, or perched position.  Legs are bent up near
     chest, arms are on knees, or folded across chest.  This behavior
     has been observed only in the younger adult female. 
     Belly scratching: Belly scratching consists of a rhythmic
     flexion of fingers or up and down movement of wrists so that the
     fingers are rubbed gently across the stomach region.  Hands may
     alternate scratching, scratch in unison, or scratch singly.  The
     animal will belly scratch while holding straw in the scratching
     hand.  The scratching posture is sitting with knees acutely bent
     and rotated outwards so the soles of feet are flat against each
     other or legs may be slightly extended with feet gripping cage
     bars.  Seen only in the older adult female.
     Stereotypic object use: Repetitive, unvarying, and seemingly
     useless manipulation of an object.
     Stereotypic noises (Auditory): Repetitive, unvarying, and
     seemingly useless production of sound.  This does not include
     the occasional production of unusual sounds or noise. 
     Motor boat sounds: Forcing air through tight lips makes a sound
     similar to a high pitched motor boat.  Emissions are of short
     duration (< 1 sec), but occur in series (one bout had 35). 
     Occurs primarily while the animal is in a resting/idle posture. 
     This behavior has only been observed in the younger silverback
     male.
     Unusual/idiosyncratic:..........

       Lip hang:     The lower lip droops loosely down exposing gums
                     and inner lips.  This behavior occurs in a
                     variety of contexts, such as after aggressive
                     encounters, during resting periods, during
                     foraging bouts, while locomoting, while
                     manipulating objects, and when head circling or
                     head swinging.  This behavior has been
                     observed only in the younger silverbacked male
                     who displays it frequently and incorporates it
                     into many behavior patterns.

 Affiliative:

   Reproductive/Sexual:

     Precopulatory:  Sexual behavior which occurs prior to
                     copulatory mounting.

     Penile erection:There is no mounting.  This is very difficult
                     to observe.

     Inspect genitalia:Close visual, tactile, or olfactory scrutiny
                     of the genitalia or genital region of another.

       Touch:        An animal briefly touches, holds, fondles, or
                     mouths the genitalia of another.

       Olfactory/sniff: An animal leans over and presumably sniffs
                     the genital region of another.  Touching
                     another's genitalia and then sniffing the
                     fingers/hand is included in this category.

     Solicit:        One animal engages in a variety of postures,
                     gestures, or movements which have a high
                     probability of eliciting sexual behavior from
                     another.

       Stare:        Female stands motionless while intensely
                     watching the male.

       Extended arm
       invitation:   Female reaches toward male with arm stretched
                     and palm facing the male.

       Head jerk:    While staring at a male, a female
                     intermittently tosses her head to one side. 
                     Her body appears relaxed and the lips are not
                     compressed.

       Present
       quadrupedal:  Female stands on all four limbs (see stand
                     quadrupedal) and orients her anogenital region
                     directly towards another animal.  She may
                     slightly elevate her hindquarters or display a
                     head jerk.

       Back into:    Female first presents and then walks backward
                     into the male.

       Open leg display:Female slides down past another while her
                     legs are spread widely apart.  This behavior
                     occurs in a playful context.

       Crouch:       While in front of another, female bends low to
                     the ground with her limbs pulled underneath
                     her.

       Ground object tap:While male is oriented towards a female, he
                     repeatedly taps the substrate with the
                     fingertips of both hands.

       Haunch scoot: While squatting, a female uses her arms to
                     propel herself towards the male.

       Quad-bounce:  While standing on all fours and facing another
                     animal, a female bounces her hindquarters up
                     and down.  Similar to a haunch bounce, but done
                     while animal is standing--may be a form of
                     masturbation as well as solicitation.
   
       Pronk:        While standing quadrupedally and facing another
                     animal, a female hops about in a jerky fashion. 
                     The hindquarters are slightly elevated.

       Pronk-mount:  A female pronks while grasping another animal
                     around the waist.

       Play present: A female presents in an "apparently playful
                     context"?.  The angle of bend at the hips is 
                     greater than in the other "present" solicit. 
                     The female may rest on her forearms and
                     occasionally look backwards (over her shoulder
                     or between her legs) at the male.

     Copulatory:

     Mount attempt:  Male attempts to make genital contact with
                     female (in any posture), but the attempt fails.

     Mount:          One animal positions itself on another such
                     that their genitalia are in contact (can be
                     same sexed), mount is of short duration with no
                     pelvic thrust observed.  Orientation of mount
                     can be dorso-ventral, ventro-ventral, or
                     ventro-lateral.

.    Copulation:     One animal mounts another in any orientation
                     (D/V, V/V, L/V).  Mounting is followed by about
                     30 to 45 sec of position adjustments before
                     pelvic thrusting begins.  During copulation the
                     female exhibits compressed lips and the male
                     exhibits pursed lips.  The female will
                     occasionally look over her shoulder at the
                     male, and sometimes wave her head side to side
                     with her eyes closed and may emit copulatory
                     whimpers.  It is very difficult to observe
                     intromission.

     Reach around:   During copulation, while in the D/V position,
                     the female twists shoulders and neck to look at
                     the male, and reaches back and touches the male
                     on the head.

     Copulatory whimpers:

     Pursed/compressed lips:

     Postcopulatory:

     Touch genitalia:Visual and tactile inspection of an
                     individual's own genitalia following
                     copulation.

     Vocalization:   After copulation.


   Play:             Repetitive, exaggerated, disjointed, and
                     seemingly nonpurposeful behavior involving two
                     or more individuals.  This behavior is
                     accompanied by the "play face" which is
                     characterized by an open mouth which partially
                     exposes the teeth, but the lips are relaxed at
                     the edges.  Social play may be accompanied by
                     soft chuckling or a throaty grumbling.  Both
                     gentle and aggressive components may be
                     observed, so it is easily confused with
                     aggression (which is generally much louder).

     Contact/direct play:Two or more individuals are engaged in play
                     involving almost continuous contact.  This
                     includes such interactive acts as:
   
                     Stand/climb on, wrestle, slap/beat, tickle,
                     poke, grab, kick, carry, ambush, tag,
                     push/nudge, swat, tap on another, object
                     tug/grab, mouth (specify act).

     No contact/
     indirect play:  Two or more individuals are involved in play
                     that does not involve direct contact.  This
                     could include such acts as:

                     Chase, throw at, ground/water slap, object
                     wave/shake/brush on ground, arm shake, chest 
                     beat, body slap (own), circle (specify act).

     Solicit play:   One animal engages in a series of postures,
                     movements, or gestures (runaway, swagger,
                     chestbeat) which has a high probability of
                     eliciting play from another.

     Parallel play:  Two animals engage in lone or object play (see
                     those categories) within proximate distance of
                     one another.

   Groom:            

     Allogroom:      One animal manipulates the fur, extremity, or
                     orifice of another.  During a grooming episode,
                     the groomer often looks intently at the portion
                     of the body which is being manipulated. 
                     Grooming may include both manual and oral
                     components.

       Manual:       Individuals use the fingers and whole hand to
                     manipulate and remove materials.

       Oral:         Use of mouth and lips to manipulate and/or
                     remove materials.

     Mutual groom:   Same as above, except the two animals groom
                     each other simultaneously.

     Allomanipulate: One animal rubs, pats at, or fondles the fur,
                     orifices, or extremities of another (not
                     including genitalia).

   Social locomotor: Involves travel while in contact with another.

     Buddywalk:      Animals walking tripedally or bipedally while
                     holding each other across the shoulder or on
                     the hips.  This form of travel could involve
                     more than one other individual.

     Carry/cling:    One animal transports or is transported by
                     another:

       Ventral carry/cling: While standing or moving quadrupedally
the focal animal transports another which is hanging on
                     to its ventrum (carry).  The walk is
                     quadrupedal or tripedal (if the focal animal is
                     using one of its arms to support the
                     passenger).  The focal animal is transported in
                     the manner described above (cling).

       Dorsal or lateral c/c:While standing or moving quadrupedally,
                     the focal animal transports another on its
                     back, side, neck, shoulders, or head (carry) or
                     is transported by another in this manner
                     (cling).

       Extended arm carry:An animal carries another (usually an
                     infant)  in the palm of the hand and walks
                     either tripedally (carrying the infant football
                     style) or quadrupedally (carrying arm is
                     inverted and weight is on the backside of the
                     hand/wrist).

       Leg drag:     An animal holds onto another's ankle and is
                     pulled along in the direction in which latter
                     animal is traveling.

   General social contact:

     Active maintenance
     of contact:     Consists of a group of actions which establish,
                     regulate, or maintain contact with another.

       Withdraw:     One animal moves from within to beyond contact
                     distance of another.

       Approach:     One animal moves from beyond contact distance
                     to within contact distance of another.

       Maintain:     Preserve social distance with a withdrawing
                     animal.

                     Follow:One animal maintains proximity (1-3 BL)
                     with another and simultaneously parallels its
                     route of travel.  Delay of follow is minimal.

                     Restrain:One animal physically prevents another
                     from moving out of contact distance.

                     Fetch/retrieve:One animal approaches and makes
                     contact (grasp, pull, push) with a withdrawing
                     animal or one who is beyond social distance (4
                     m), which results in that individual returning
                     to an original location or group.

                     Herd:An individual guides the movement of an
                     individual or troop by a series of approaches
                     and withdrawals.

       Greet:        Involves a group of actions which occur when
                     individuals are seeing each other for the first
                     time or are entering into the contact distance
                     of one another.  These actions are typically
                     brief (momentary to 5 sec).

                     Muzzle-muzzle:One animal places its face within
                     15 cm of another's muzzle.
     
                     Kiss:One animal places its lips against the
                     head or face of another.
   
                     Touch:One animal reaches with any limb and
                     makes contact with another on any part of the
                     body except the genital region.  The contact is
                     momentary (1-2 sec).
 
                     Embrace: One animal wraps arms and/or legs
                     around the body of another.  Duration is short
                     (< 10 sec).

                     Grasp:One animal momentarily seizes another
                     with the hand(s).

                     Offer food:An animal holds food in the hand and
                     then extends that hand towards another.

     Inactive social contact:Individuals are inactive, are within
                     social proximity (0-4 m) of one another, and
                     have the option to withdraw from social
                     proximity.

       Huddle:       Two or more individuals are inactive with
                     torsos in direct contact, lateral or ventral. 
                     Arms may be wrapped around one another.

       Rest in contact:Individuals are inactive with some part of
                     the body (exclusive of the position described
                     in huddle) in contact i.e., back to back, toes
                     touching, arms intertwined.

       Rest proximate:Individuals are inactive, not in contact, but
                     within arms length of one another.

       Rest distal:  Individuals are inactive, not in contact,
                     greater than arms length away but within 4 m of
                     one another.

 Agonistic:

   Submissive/avoidance:

     Turn away:      An animal moves its body so that it is oriented
                     away from another, but does not travel.

     Avoid:          An animal moves out of the path of an
                     approaching animal or takes a less direct route
                     around that animal.

     Hide:           An animal removes itself from the view of
                     another.  This may occur following an
                     aggressive attack or threat.

     Crouch/crawl:   This behavior can take two forms: 1) An animal
                     bends all four limbs, presses its ventrum to
                     the ground, and may try to travel while in this
                     position; or 2) the animal may crouch while in
                     one of the sitting positions by lowering the
                     head, hunching the shoulders, and often
                     covering the head with an arm.

     Present:        May take two forms.  One form is similar to a
                     sexual present, but is much briefer.  It may be
                     accompanied by a series of brief glances
                     directed towards the presentee.  An animal may 
                     also "offer-up" or present an arm to the
                     presentee. (specify act)

     Run away:       An animal moves rapidly (with a running gait)
                     away from another.

   Supplant:         

     Supplant position:An animal approaches another which then moves
                     away. The former assumes the position of the
                     latter.

     Supplant take object:An animal takes an object from another or
                     picks up one which was dropped as the former
                     approached.

   Threat:           Posture, facial expression, or movement,
                     usually accompanied by a vocalization, which
                     results in keeping rivals at a distance,
                     obtaining desired objects or space, and
                     controlling another's behavior without undue
                     expenditure of energy or risk of injury
                     (McFarland, 1982, slightly modified).

     Threat:         May appear to be undirected and for our
                     purposes, does not involve movement towards
                     another.

       Stare:        An animal looks towards and fixates on another
                     for an extended time.  The brow is furrowed and
                     the face is tense.

       Tight lipped face:An animal's lips are compressed tightly so
                     that the upper and lower lips protruded
                     somewhat.

       Rigid quadrupedal
       stance:       An animal assumes a quadrupedal stance, with
                     the legs widely spaced and placed behind the
                     torso.  The arms are rigid with elbows rotated
                     outwards, and the weight of the upper body on
                     the knuckles.

       Rigid quadrupedal
       walk:         From the rigid quadrupedal stand, an animal
                     moves forwards with short stiff steps.  The
                     walking speed may be faster than normal,
                     creating a trot like gait.  This is undirected.

       Object slap:  An animal slaps ground, rock, wall, tree, door,
                     etc., with the palm or back of the hand.  This
                     usually results in a loud sound.

       Head divert:  One animal turns its head away from another in
                     an exaggerated manner and glances repeatedly
                     and rapidly back at them.

       Yawn:         The mouth opens widely, head tips back, lips 
                     are pulled back so that the teeth are exposed. 
                     Looks like exaggerated form of a yawn.

       Chest beat:   An animal repetitively strikes the chest or
                     belly area with outstretched or cupped hands. 
                     The animal is usually bipedal; either walking,
                     running, standing, or standing and rapidly
                     swaying.  An animal may also strike his/her
                     chest once with a cupped hand.  This can occur
                     in any posture.  The chest beat, like all
                     agonistic behavior, may include a vocal
                     component such as a roar or pant hoots.  Note:
                     chest beating can be observed in a number of
                     contexts: i.e., lone and social play,
                     reproductive, and agonistic.

       Ground slap:  An animal thumps the ground with one or both
                     hands.  The strike may be with open hands, back
                     of hands, or side of closed fists.

       Directed chest beat:An animal chest beats while oriented
                     towards another.

       Directed object slap:An animal strikes at an object with an
                     open hand while oriented towards another.

       Open mouth threat:An animal looks at another and lunges
                     forward with a tense, open-mouthed expression. 
                     The lips are raised and pulled back so that the
                     canines are exposed; muscles in the forehead
                     are taut.

       Threat display:A series of threats, postures, facial
                     expressions, or movements, that are accompanied
                     by a roar, bar, or rumble vocalization.  Many
                     of the threat displays include a rapid advance
                     directed towards specific individuals.
       Bluff charge: An animal runs quadrupedally on the diagonal
                     past another but does not make contact.  The
                     bluff charge usually ends in a slide, and may
                     be accompanied by an object flail, drag, or
                     throw.

       Rush charge:  An animal runs quadrupedally towards, but stops
                     just short of another.

       Object grab:  One animal grasps or snatches an object (food,
                     browse, or otherwise) from another.

     Attack:         Includes hostile acts involving active pursuit
                     or physical contact between two individuals.

       Hit:          While rapidly charging towards another
                     (bipedally or quadrupedally), one animal
                     reaches out and strikes the other with an open
hand or fist.
 
       Hit with object:An animal uses an object to strike out and
                     makes contact with another.  This behavior
                     occurs at the end of a charge.

       Hit/slap:     An animal strikes another with its hand(s), but
                     there is not charge component to this behavior.

       Push:         Animal uses arms or legs to forcefully move
                     another away.

       Pull:         An animal grasps another and forcefully moves
                     them closer.

       Chase:        An animal runs in pursuit of another which is
                     rapidly withdrawing.

       Bite:         An animal closes its mouth on the extremities
                     or body of another.  Damage may or may not
                     occur.








  Author Note:
The authors would like to acknowledge that this ethogram was derived
in part from ethograms provided by the Los Angeles Zoo, Zoo Atlanta,
and Seattle, as well as through observations of the gorillas located
at the Dallas Zoo.
.Bowen, R. A.  (1980).   The behaviour of three hand-reared lowland
                         gorillas (Gorilla gorilla gorilla) with
                         emphasis on the response to a change in
                         accommodation.  Dodo, R. Jersey Wildlife
                         Preservation Trust, 17, 63-79.

  Purpose:           To measure effects of change in accommodation.

  Sampling methods:  Scans, one minute intervals for one hour
                     periods.

  Ethogram key words:Activity budgets/General behaviors, Environment
change, Influence of.

  Behaviors:         (full listing available from author)

  Resting:           Lying, sitting, standing, clinging and hanging,
                     except when accompanied by any other behavior;
                     thus leaving only those occasions on which the
                     animal was sedentary in both gross and local
                     terms (apart from visual behavior or gestures).

  Gross locomotion:  Walking, running, climbing, brachiating,
                     swinging, rolling, and wrestling.

  Local manipulation:Manipulating with hands or feet, mouthing,
                     scratching, hairplucking, grooming,
                     nose-picking, coprophagy, drinking water or
                     urine, foraging, feeding, and mauling (gentle
                     wrestling).

  Social play:       Wrestling, mauling, sparring, chasing and
                     social grooming.

  Self-directed:     Grooming, scratching, plucking hairs and
                     nose-picking.

  Visual:            Watching, looking at, and glancing at one
                     another.

  Gestures:          Including facial expressions (play face and
                     fear grin), playing-walking/running, tapping
                     and slapping objects/substrate, chest beating
                     and patting parts of the body.

  Public-oriented:   Watching or interacting with members of the
                     public, by throwing items of food or play
                     objects, for example.

  Feeding:           Eating, drinking, foraging for food, coprophagy
                     and drinking urine.

  Stereotyped:       'Rocking' and 'shuffling.'  These are
                     behavioural abnormalities peculiar to captivity
                     in which the animal sits, sometimes holding a
                     quantity of wood-wool or similar nesting
                     material, and either rocks back and forth with
                     it in a sort of scrubbing motion,  or shuffles
                     backwards with it, using the hands placed on
                     the surface in front as a means of propulsion.
.Brown, S. G.  (1988).  Play behaviour in lowland gorillas: Age
                         differences, sex differences, and possible
                         functions.  Primates, 29(2):219-228.

  Purpose:           To document play in a large sample of gorilla
                     groups, in order to better understand the
                     possible functions of social play in gorillas.

  Sampling:          Frequencies recorded.

  Ethogram key words:Social behavior.

  Behaviors:

  Solitary play:     
    
    Rotational/locomotor:

                     Twirl:Spinning around bipedally.

                     Somersault:Turning the body one full rotation
                     head over heels.

                     Roll:Turning the body from side to side while
                     supine; can be complete or incomplete rotation.

                     Run:Loose limbed gait, usually bipedal.

                     Self:Slapping the face, clapping hands, or
                     rubbing oneself without grooming.

  Object play:       Play with inanimate objects and other species.

    Climb/swing:     From a tree, boulder, or ledge.

    Manipulate:      An external object such as grass, dirt, or a
                     branch.

    Throw:           An external object while in a relaxed stance.

    Swing:           An external object while in a relaxed stance.

    Water:           Water is used in the play sequence.

  Social play:       

    Moderate play:

                     Open-mouth-
                     wrestling:Limbs entwined while sitting or
                     laying; gorillas roll/twist together placing
                     open mouths on each other.
    Active play:     

                     Rough/tumble:Vigorous bipedal wrestling.

                     Chasing:One gorilla rapidly following another,
                     both with relaxed running gaits.
  Chest-pounding
.duBois, T., Mead, J., & Cox, C.  Lowland gorillas: Ethogram.  Los
                                   Angeles Zoo.

  Ethogram key words:Activity budgets/General behaviors, Social
behavior, Sexual behavior, Display/Agonistic behavior.
Abnormal/Stereotypic behavior, Development, Maternal behavior,
Vocalizations.

  Behaviors:

  Masturbation:

                     Manual:An animal rubs its genitals repeatedly
                     with hands.

                     Pelvic rub:A female backs up to an object
                     (tree, rock, etc.) and rubs anogenital area
                     against it.

                     Haunch-bounce:While in a squatting position, a
                     female bounces up and down on her haunches. 
                     (Referred to as Evie's "hiccuping.")

                     Coo vocalization:Pursed lip vocalization
                     sometimes given when female's masturbate.

  Sexual behavior:

                     Anogenital touch:An animal examines or fondles
                     the anogenital region of another.

                     Anogenital sniff:An animal sniffs the
                     anogenital region of another, excluding
                     occasions when the latter animal has just
                     defecated or has feces on fur.

                     Stare:A female stands motionless while
                     intensely observing the male.

                     Head jerk:While staring, a female
                     intermittently jerks head to the side.  The
                     body is relaxed and lips are not compressed. 
                     (Referred to as "the come hither look".)

                     Extended arm invitation:A female extends her
                     arm with open palm toward the male.

                     Ground or object tap:While orienting toward the
                     male, a female repeatedly taps ground, rock or
                     log with both hands (as if playing a drum).

                     Haunch-scoot:While squatting, a female uses her
                     arms to propel herself on her haunches toward
                     the male.

                     Quadrupedal-bounce:While standing on all fours
                     and facing another animal, a female bounces
                     hindquarters up and down.  (Similar to
                     haunch-bounce but done while animal is standing
                     - may be form of masturbation as well as sexual
                     solicitation.)

                     Pronk:While standing on all fours and facing
                     another  animal, a female hops about in a jerky
                     fashion.  The hindquarters tend to be slightly
                     elevated.  (Behavior "invented" by Cleo.)

                     Pronk-mount:A female pronks while grasping
                     another animal around the waist.

                     Present:A female stands quadrupedally and
                     orients anogenital region directly toward
                     another animal; hindquarters are slightly
                     elevated (but not an exaggerated display as in
                     some primates).

                     Back-into:While presenting, a female walks
                     backwards directly into another animal.

                     Play-present:A female presents in a playful
                     context.  Form is generally exaggerated, e.g.,
                     animal bends way over with head close to ground
                     and rump in the air. (May occur only in young
                     animals.)
                     Spread-legs display:A female slides on an
                     incline past another with legs spread widely
                     apart.  Occurs in a playful context.

                     Crouch:A female bends low to the ground with
                     her limbs pulled in.  Another animal may then
                     mount dorso-ventrally.

                     Lie prone:An animal lies flat on the ground
                     face downward during a sexual encounter.

                     Lie supine:An animal lies flat on the ground
                     face upward during a sexual encounter.

                     Dorso-ventral mount:An animal mounts another
                     from behind.

                     Ventro-ventral mount:An animal mounts another
                     from the front.  (Both animals face each
                     other.)

                     Ventro-ventral mount
                     of infant:An adult animal stands quadrupedally
                     over a supine infant and thrusts without making
                     contact.

                     Awkward mount:An animal mounts another, but
                     orientation is improper, e.g., mounts from
                     side, etc.

                     Mount and thrust:An animal mounts another and
thrusts, but intromission does not occur.

                     Copulation:An animal mounts another and
                     intromission occurs (hard to determine whether
                     ejaculation occurs).

                     Coo vocalization:Pursed lips vocalization given
                     by female which may occur before, during or
                     after copulation.

                     Reach and touch:During copulation, a female
                     reaches around with one arm and touches male on
                     the head.

. Social play:       

                     Social play:Repetitive, exaggerated,
                     disjointed, and seemingly non-purposeful
                     behavior involving two or more animals.  May
                     have both gentle and aggressive components. 
                     The list is almost infinite but includes the
                     following:

                     Climb or stand on another, turn circles around
                     another, piggy back ride, peek-a-boo, mouth,
                     tickle, foot dangle (an animal dangles foot in
                     front of another), tap on another, dangle (an
                     animal holds an infant by its hands and allows
                     it to dangle in the air), arm shake, run-away
                     (one animal runs away from another as an
                     invitation to chase), chase or play tag,
                     swagger walk, chest beat, beat or slap other
                     body part, ground or water slap, object wave,
                     shake, or brush on ground (usually a branch),
                     object throw/hit, object tug/grab, swat, push
                     or nudge, grab/pull, kick, wrestle, bite,
                     play-face (relaxed, open-mouth expression with
                     teeth exposed).

  Social other/maternal:

                     Approach:An animal draws closer to another:
                     because of context and relaxed body posture,
                     motivation presumed to be friendly.
                     
                     Follow:An animal walks behind another; because
                     of context and relaxed body posture, motivation
                     presumed to be friendly.

                     Muzzle-muzzle:An animal places its face
                     directly in front of another (within 6 inches).

                     Groom/inspect:Using the hands or mouth, an
                     animal grooms or examines another's body parts,
                     excluding the anogenital region except when
                     feces being removed.

                     Active contact:Friendly contact behaviors which
                     occur outside of play context.  Includes the
                     following:
                     
                     Touch (excluding the genitals), grasp, sniff
                     (an animal sniffs another's body parts,
                     excluding anogenital region unless feces are
                     obviously being sniffed), mouth, kiss (an
                     animal puckers its lips against head or face of
                     another), pat or caress, hug or embrace, hold,
                     huddle (animals sit in groups of two or more
                     with bodies in close contact).

                     Passive contact:An animal sits, lies, or
                     sometimes walks with body in contact with
                     another.  Behavior is not as "intimate" as
                     active contact, e.g., animals may sit with only
                     feet touching. 
 
                     Inside with:Two or more animals, excluding
                     infants being carried, are inside cave or
                     planter together and are not visible to the
                     observer; assume association to be affiliative,
                     although specific behaviors not visible.

                     Food offer:An animal offers food, browse, or
                     regurgitate to another.

  Special social behaviors observed in mothers or caretakers:

                     Rock:A mother supports her infant in arms or on
                     lap and rocks it back and forth.

                     Jiggle-bounce:A mother supports her infant in
                     lap or on legs and bounces and jiggles the
                     infant.

                     Dangle-groom:A mother dangles her infant by one
ankle and grooms the anal area.

                     Flex-manipulate:An animal bends and manipulates
                     the infant's body parts.

                     Coax:A mother places her infant on the ground
                     and then walks away and stands or sits at a
                     distance.  The mother observes infant
                     intensely, seemingly encouraging it to
                     approach.

                     Retrieve:An animal picks up an infant when
                     infant is alone.

                     Restrain:An animal holds an infant firmly when
                     the infant attempts to leave.

                     Shield:An animal uses its arms or body to
                     shield an infant from another animal or other
                     potential danger.

  Special social behaviors observed in infants:

                     Climb on or over:An infant climbs on or over
                     another animal.
                     
                     Nose/eye poke:An infant pokes its finger in
                     another's nose or eye.

                     Manipulate mother's
                     nipples:An infant pulls at or handles mother's
                     nipples (not to be confused with rooting
                     behavior).

                     Distress vocalization:An infant whines or cries
                     when alone; appears to be an attempt to get
                     mother to return.

  Agonistic:

                     Turn away:An animal turns away from another in
                     order to avoid, but does not vacate its spot.
 
                     Avoid:An animal moves away from or gets out of
                     path of another.

                     Crouch:An animal crouches when threatened or
                     attacked by another.

                     Hide:An animal presses itself against a surface
                     so that it is hidden or partially hidden from
                     the view of another.

                     Supplant:One animal approaches another and the
                     latter moves away.  The approaching animal then
                     takes that spot.

                     Stare:A fixed, unwavering stare at another with
                     brow furrowed and facial muscles tense.

                     Tight-lip face:A facial expression in which the
                     lips are tightly compressed and the head
                     diverges from side to side.

                     Stiff stance:An animal stands with the legs
                     held rigidly and the body stiff and erect;
                     tight lip face often accompanies.

                     Strut-walk:While in the stiff stance, an animal
                     walks with stiff, short steps.

                     Directed chest beat:An animal chest beats while
                     orienting toward another gorilla in the same
                     exhibit.

                     Directed object slap:An animal orients toward
                     another and slaps ground, rock, wall, inside
                     door, or makes loud splash in the pond.

                     Bluff charge:An animal runs on the diagonal
                     past another but does not make contact.

                     Rush charge:An animal rushes up to and stops
                     just short of another.

                     Charge with chest beat:While charging, an
                     animal beats its chest.

                     Charge w/object display:While charging, an
                     animal waves or throws an object or slaps
                     ground with an object (usually browse).

                     Charge and hit:While charging, an animal hits
                     another with an object (usually browse).

                     Leg kick:An animal kicks its leg in the air at
                     the end of a charge, chest beat or object
                     display.

                     Object grab:An animal snatches an object or
                     browse away from another.

                     Object drop or throw:An animal drops an object
                     down on another from  above, or throws an
                     object at another.

                     Hit with object:An animal hits another with an
                     object.

                     Hit/slap:With hands.

                     Push

                     Grab/pull

                     Chase

                     Open-mouth threat:A tense, open-mouth
                     expression with lips raised and pulled back so
                     that canines are exposed; muscles and the
                     forehead are taut and drawn back.

  Agonistic -- Vocalizations:

                     Soft pant-hoot:A quiet vocalization in which an
                     animal makes rapid exhalations through "o"
                     shaped lips.  The abdomen may be seen heaving
                     up and down.

                     Scream

                     Pig grunts:Short, rough, guttural grunts.

                     Growl:A deep, guttural noise similar to a dog's
                     growl.

                     Bark:A deep, gruff noise similar to a dog's
                     bark.

                     Roar:A loud explosive outburst given by a male
                     silverback.

  Herd/herded:

                     Herd:A silverback male controls the movement of
                     a group by following at a close distance
                     (formerly called pursue; may be an attempt to
                     keep females together and in sight.)

                     Herded:Refers to a group being herded by a
                     silverback male.  Animals appear to be avoiding
                     the male as they move away whenever he
                     approaches.

  Visual attention:

                     Observe:An animal watches another gorilla in
                     the same exhibit.
                     
To outside gorillas:     [The following behaviors are directed to
gorillas in 1)an adjacent exhibit, 2) in night quarters, or to 3)
"missing" gorillas (gorillas which have recently been sent out).]

                     Observe:An animal watches another gorilla in an
                     adjacent exhibit.

                     Display:Includes behaviors such as chest
                     beating, object throwing, waving, etc., which
                     are directed toward gorillas in an adjacent
                     exhibit.

                     Bang door:An animal bangs on the inside door
                     when there are group members kept inside night
                     quarters.

                     Search:An animal travels about exhibit or goes
                     in and out of cave seemingly looking for a
                     missing group member.

                     Hoot series vocalization:Low pitched "hoo hoo
                     hoos" which can build up to more plaintive
                     sounding, higher-pitched "hoos."  Vocalization
                     often terminated by banging on door and has
                     been heard only when group members are missing.

  Stereotyped behavior:

                     Head toss:An animal repeatedly tosses its head
                     in circular fashion; sometimes followed by
                     regurgitation and reingestion.

                     Arm fling:While sitting, an animal repeatedly
flings its arm out for no obvious reason.

                     Hair pluck:An animal plucks hair from its own
                     body, usually the facial region.

                     Self-flagellation:While sitting, an animal
                     flogs itself with browse.
Regurgitation and reingestion:   [Refers to behaviors in which food
is brought up from the stomach to the mouth and then eaten. 
Gorillas tend to use idiosyncratic methods.]
                     
                     Reg. into cupped hand:An animal regurgitates
                     into hand and then eats out of hand.

                     Reg. onto substrate:An animal bends over,
                     regurgitates onto ground and then ingests
                     directly from ground.

                     Bent-arm regurgitate:An animal lifts and bends
                     arm at elbow while regurgitating in a jerky
                     fashion into cupped hand.

                     Suck-regurgitate:An animal brings lips into
                     extreme sucking position and brings regurgitate
                     up into mouth.

                     Backward neckAn animal leans head backward with
                     neck stretched taut,
                     stretch regurgitate: then regurgitates with
                     mouth closed.

                     Regurgitate into mouth:An animal makes "o"
                     shape with lips and brings regurgitate up into
                     mouth.

                     Head jerk regurgitate:An animal jerks head in
                     circular fashion while  bringing up
                     regurgitate.

                     Bend over regurgitate:While sitting, an animal
                     bends over with mouth close to the ground and
                     regurgitates into mouth.

  Self-play and object explore:

                     Self-play:Includes all solitary play behaviors
                     which are not directed to other animals or to
                     the zoo patrons, including:

                     Rock body, roll on ground or grass, somersault,
                     turn circles, thigh slap/pat, chest pat (very
                     mild chest beating), lip flap (using fingers),
                     clap hands, manipulate body parts (exaggerated
                     body or head rubbing, pulling on toes, etc.),
                     water play (wading, splashing, etc.), object
                     play (waving branches, tossing leaving in air,
                     playing with tubs, burlaps sacks, etc.).

                     Object explore and Refers to all behaviors in
                     which an animal uses its hands
                     manipulate:to investigate or handle objects,
                     excluding those which are preparatory to
                     feeding, including:

                     Browse manipulate, browse or object wear (an
                     animal drapes browse, burlap sack, etc., over
                     back, shoulders, head), other vegetation
                     inspect/manipulate (rubbing grass on body,
                     throwing grass clumps in air, etc.),
                     manipulate feces or regurgitate (not associated
                     with ingestion), poke at dirt/dig holes, rub
                     self with dirt, mouth objects (including
                     biting, sucking, licking, chewing, or
                     manipulating with tongue), manipulate
                     miscellaneous objects (such as drain covers),
                     items thrown in by patrons, etc.), observe
                     objects (visual examination of an object which
                     is being held or which is within reach), sniff
                     objects, and carry objects (in mouth, hand, or
                     on small of back).

  Food getting and ingestion:

                     Ingest:Includes the following: eat browse, eat
                     grass, weeds, leaves, bark, flowers, or other
                     vegetation, eat miscellaneous food items
                     (including food thrown in by keepers and
                     patrons), spit food out, drink, eat food out of
                     another animal's mouth (seen in infants).

                     Forage:Behaviors which are preparatory to
                     feeding.  Includes the following: gathering,
                     carrying, or dragging browse, bark stripping,
                     gathering and eating seeds, picking and/or
                     carrying grass, weeds, leaves, bark, flowers,
                     or other  vegetation or food items.

  Audience interaction:

                     Audience interaction:Includes all behaviors in
                     which an animal interacts with humans (keepers,
                     patrons, etc.), or vehicles drawn in front of
                     an exhibit.  Includes: approach/observe,
                     extend arm (animal stretches arm outward toward
                     audience, probably begging for food), point at,
                     clap hands, arm shake, blow kisses, self-play
                     behaviors (which are obviously directed to
                     audience), chest pats, splash water, object
                     throw (including feces, dirt clods, rocks,
                     etc.), threaten audience (animal directs
                     agonistic behavior toward humans, including
                     growl, roar, open-mouth threat, bluff charge,
                     chest beat).

  Carry/cling or ride:

                     Ventral carry/cling:While standing or moving
                     quadrupedally an animal carries an infant who
                     clings to the ventrum. 

                     Dorsal or lateral
                     carry/cling:While standing or moving
                     quadrupedally an animal carries an infant on
                     the back, side, neck, shoulders or head.

                     Leg drag:An infant grasps another's ankle and
                     clings while the latter animal walks about.
                     
                     Extended arm carry:An animal carries an infant
                     in palm of outstretched hand and walks
                     tripedally (the "football carry").

                     Tripedal walk:An animal holds infant against
                     the ventrum and walks tripedally.

Locomotion:    [Includes all forms of movement and travel occurring
outside of a friendly/social or agonistic context.  Also excludes
self play and occasions when an animal is carrying an infant.]

                     Walk quadrupedally:An animal walks on four
                     legs.

                     Walk tripedally:An animal walks on three legs.

                     Walk bipedally:An animal walks on two legs.

                     Stand quadrupedally

                     Stand bipedally

                     Climb

                     Scoot

                     Slide

                     Roll

                     Jump

                     Hang by arms

                     Swing by arms

                     Run: (in non-agonistic context).

                     Change stance:Go from sitting to standing
                     position, etc.

  Special locomotor behaviors observed in infants:

                     Arm/leg flail:An infant jerks its arms and/or
                     legs about in a seemingly random fashion.

                     Root:An infant gropes and moves head about
                     mother's ventrum in search of mother's nipple.

                     Crawl:An infant drags its prone body along the
                     ground.

                     Toddle:An infant walks quadrupedally with
                     short, uncertain steps.

  Rest and body maintenance:

                     Rest:Periods of quiet repose in which any of
                     the following behaviors may occur:

                     Sit/lie without performing other behaviors,
                     sleep, yawn, stretch, cough, comfort movements
                     (slight position shifts or rolling over while
                     resting).

                     Self-exploration and
                     body maintenance:Includes the following
                     behaviors:

                     Groom/self-inspect (an animal picks at or
                     examines its toes, hands, or any other body
                     parts), oral explore (an animal mouths or licks
                     its body parts), scratch, nose-pick, sniff
                     self, rub eyes, pick teeth, rub self, poke
                     anus/remove feces.

                     Urinate/defecate

                     Vomit:(when not part of R/R).

                     Look out/look about:An animal sits or lies
                     quietly but appears to be observing
                     environment, although specific object of focus
                     is not apparent (same as "stationary alert").

  Special rest behaviors seen in infants:
 
                     Nurse:An infant suckles at its mother's breast
                     (it is often hard for an observer to tell
                     whether an infant is nursing or sleeping, so we
                     are including "nurse" under rest behaviors).

Other:   [These include behaviors which do not readily fall into any
category or whose functions are unclear.]

                     Non-directed agonistic:Includes agonistic
                     behaviors such as chest beat, growl, object
                     slap, etc., which do not appear directed toward
                     other gorillas or to audience, and which occur
                     outside of play context.

                     Interspecific agonism:An animal directs
                     agonistic behavior toward animals other than
                     gorillas or humans (e.g., swiping at insects,
                     chasing peacocks, etc.).
                     Visual exploreIncludes the following: visual
                     attention toward objects in
                     environment:environment which are not within
                     reach (such as planes, helicopters, etc.), and
                     observe animals other than gorillas or humans
                     (such as peacocks, insects, etc.).

                     Eat feces

                     Drink urine

                     Eat another's regurgitate

                     Monitor/search:An animal looks about exhibit as
                     if to locate the whereabouts of other group
                     members.

                     Armpit sniff/touch/
                     manipulate:An animal sniffs or touches own
                     armpit, or manipulates area under armpit.

                     Nipple rub:An animal repeatedly rubs its own
                     nipple.

                     Self-directed lactating
                     behaviors:Behaviors shown by lactating mothers
                     including:

                     Manipulate nipples, squirt milk from nipples,
                     and bend over to drink milk from own nipples.

                     Escape/escape attempt:An animal attempts or
                     succeeds in climbing out of exhibit.  May stand
                     on back of another gorilla to reach handhold.

                     Nest-build:An animal manipulates browse around
                     itself to form a crude, circular nest.

                     Not visible:An animal is not visible to the
                     observer excluding cases of inside with..
Elliott, R. C.  (1976).  Observations on a small group of mountain
                         gorillas (Gorilla gorilla beringei).  Folia
                         Primatologica, 25, 12-24.

  Purpose:           To summarize the characteristics of range and
                     behaviour in a small gorilla group. 

  Sampling methods:  Modified time sampling technique to record
                     inter-individual distance data.  Position
                     mapped at 10 min intervals.  Longhand field
                     notes apparently taken on behavior, but no info
                     on how or definitions.

  Ethogram key words:Activity budgets/General behaviors, Field study,
Gorilla g. beringei. 
  
  Behaviors:

  Agonistic behavior
  Allogrooming
  Social play
  Vocalizations
.Fischer, R. B.  (1984).  Observations of group introductions in
                         lowland gorillas.  Behavioural Processes,
                         9, 293-296.

  Purpose:           To investigate agonistic responses during and
                     just after group formation.

  Sampling:          Two hours of continuous observation followed
                     each introduction. 

  Ethogram key words:Display/Agonistic behavior,
Introductions/Socialization.

  Behaviors:         

  Nonagonistic behaviors:

                     Proximity:When two or more animals were less
                     than 4 m apart for at least one minute. 
                     
                     Arm-over:Lateral extension of arm towards
                     another.  Arm was typically relaxed. 
                     Aggression has not been observed following
                     approach preceded by arm-over.

  Agonistic behaviors:

                     Contact aggression:Frequencies of hitting,
                     biting or grappling observed between
                     individuals.

                     Noncontact aggression:Instances of directed
                     pursuit (subsuming charging and chasing),
                     demonstration (species typical postural and
                     facial displays, chest beating, throwing, or
                     aggressively striking an object as part of a
                     sequence of activities), and lunging. 
                     Behaviors not clearly directed toward a
                     specific individual were omitted.
.Fossey, D.  (1972).  Vocalizations of the mountain gorilla (Gorilla
                    gorilla beringei).  Animal Behaviour, 20:36-53.

  Purpose:           To describe the vocalizations of the mountain
                     gorilla.

  Ethogram key words:Vocalizations, Field study, Gorilla g. beringei.

  Behaviors:         (See also: Fossey, D. (1983).  Gorillas in the
                     mist.  Boston: Houghton Mifflin.)

  Roar:              Monosyllabic loud outburst of low-pitched harsh
                     sound, lasting from .20 to .65 seconds,
                     beginning and ending abruptly.  Individual
                     differences in frequency concentrations.  Heard
                     only from silverbacks in situations of stress
                     or threat, and primarily directed at human
                     beings, although occasionally at buffalo herds. 
                     Always followed, on the part of the emitter,
                     with varying degrees of display, ranging from
                     bluff charges to small forward lunges.

  Alarm calls:       

                     Scream:Shrill and prolonged emission of
                     extremely loud sound, lasting up to 2.13
                     seconds and repeated as often as 10 times. 
                     Individual differences not denoted.  Screams
                     heard from all age and sex classes, but most
                     frequently from silverbacks.  Vocalization
                     heard most often during intragroup disputes,
                     though could be directed toward human beings or
                     ravens if alarm rather than threat was
                     motivation for call.

                     Wraagh:Explosive monosyllabic loud vocal
                     outburst not as deep as a roar nor as shrill as
                     a scream.  Began and ended abruptly and lasted
                     between .2 and .8 second.  Individual
                     differences in sound, which were more
                     harmonically structured than roars.  Heard from
                     all adults but most frequently from
                     silverbacks.  Usually precipitated by sudden
                     situations of stress--the unexpected arrival of
                     an observer, etc.  Most effective in scattering
                     group members and never accompanied by
                     aggressive display behavior.

                     Question bark:Characteristic composition of
                     three notes with the first and third lower than
                     the middle, as if asking the question "Who are
                     you?"  Sound was short, lasting between .2 and
                     .3 second, heard more from silverbacks. 
                     Usually in situations of mild alarm or
                     curiosity and was common response to discovery
                     of obscured observer, etc.

                     Cries:Resembling wails of human infants, could
                     build up into shrieks much like human beings'
                     temper tantrums.  Emitted between .03 and .05
                     seconds apart, and could last for nearly 19
                     seconds.  Wails had four distinct frequency
                     concentrations, but shrieks were much less
                     structured.  Cries were heard only from infants
                     or young juveniles, frequently when left alone. 
                     Cries built up into temper tantrums if
                     stressful situation was prolonged.

. Coordination Vocalizations:

                     Pig-grunts:Series of short, rough, guttural
                     noises, pig-grunts are usually delivered
                     between .15 and .4 seconds apart in sequences
                     of nine or ten outbursts.  Resembling grunting
                     of pigs feeding, tended to become louder and
                     more closely spaced if prolonged.  Most
                     frequently heard during traveling, when trail
                     disputes and altercations over limited food
                     resources were more apt to occur.  Pig-grunts
                     were effective rebuttal vocalizations and
                     disciplinary enforcements between adults and
                     young.

                     Belch vocalizations:Deep, prolonged rumbles
                     (naoom, naoom, naoom) rather like
                     throat-clearing utterances.  The belch
                     vocalization is one of the most complex because
                     of multiple intergradations and functional
                     variations.  Variations heard from all sex and
                     age classes.  For expressing contentment and
                     verifying position; a slightly shortened belch
                     vocalization was often used when mildly
                     disciplining gorilla young or observers. 

                     Chuckles:Raspy expirations of noise verge in
                     intensity depending on degree of play involved. 
                     Irregularly spaced spurts of sound varying from
                     .02 to .1 seconds with low frequency
                     concentration.  No individual differences.

  Intergroup vocalizations:

                     Hootseries preceding
                     chestbeats:Given with or without a terminating
                     chestbeat, consists of prolonged distinct
                     hoo-hoo-hoos.  Low-pitched, often undetectable
                     to human ear at beginning, but usually built up
                     into plaintive-sounding and longer hoots
                     toward the end.  The lengthier the series, the
                     more fluctuations in harmony and phasing. 
                     Frequencies ranged between 1.4 and 1.8
                     kilocycles per second for as many as
                     eighty-four hoots per second.  Silverbacks were
                     most frequent emitters.  Depending on distance
                     between groups, would or would not terminate
                     vocalizations with displays. 

.Fossey, D.  (1979).  Development of the mountain gorilla (Gorilla g.
                    beringei): The first thirty-six months.  In D.
                    A. Hamburg and E. R. McCown (Eds.), Great apes
                    of Africa (pp. 139-186).  Menlo Park, CA:
                    Benjamin-Cummings.

  Purpose:           To begin describing quantitatively individual
                     differences of mother-infant relations; also
                     describes behavior during pregnancy.

  Sampling method:   Ad lib field notes, later classified under
                     specific subject headings.  Data were then
                     grouped according to two month units of infant
                     age (e.g., 1-2 months old).  This paper also
                     describes physical development in detail.

  Ethogram key words:Development, Pregnancy, Behavior during, Field
study, Gorilla g. beringei.

  Behaviors:         (3 main categories: maintenance activities,
                     maternal interactions, and social awareness,
                     listed according to time unit.) (See also:
                     Fossey, D. (1983).  Gorillas in the mist. 
                     Boston: Houghton Mifflin.)


  Newborn (first 24 hrs):

                     Whines:Weak, puppy-type.
                     Grasping reflex
                     Rooting
                     Nuzzling of nipple
                     Sleep
                     Ventral carry
                     Grooming by mother
                     Nurse

  1-2 months:

                     Ventral clinging
                     Ventral carry
                     Low abdominal ride
                     Thigh ride
                     Dorsal ride
                     Rooting
                     Nuzzling of nipple
                     Mouthing or chewing of
                       vegetation debris
                     Sleep
                     Whines
                     Wails:Loud, high pitched.
                     Exploring own body
                     Rolling over
                     Creeping
                     Look around
                     Grooming by mother
                     Nurse

. 2-4 months:

                     Ventral clinging
                     Ventral carry
                     Dorsal ride
                     Crawl
                     Sitting attempts
                     Kick, whack, push
                     Exploratory play,
                       surrounding
                       vegetation:Gross, jerky extensions with
                     fingers widely spread before object is
                     contacted, usually only grasp object.
                     Exploratory play,
                       mother's body:Unbalanced crawling, sliding,
patting, and hair pulling.
                     Discipline:Mock-biting, pushing back at infant.
                     Eating vegetation debris
                     Grooming by mother
                     Distress face
                     Play face
                     Whines
                     Wails
                     Screeches
                     Panting play chuckles
                     Nurse

  4-6 months:

                     Ventral carry
                     Ventral cling
                     Dorsal ride
                     Grooming by mother
                     Nurse
                     Exploratory play,
                       surrounding
                       vegetation:Includes manipulation of
                     vegetation.
                     Exploratory play,
                       mother's body:More strenuous, includes
                     mock-wrestling with extremities.
                     Solo play:Patting, clapping and whacking own
                     body.
                     Restrain infant
                     Eating vegetation:No food prep, just plucking
                     leaves or gnawing.
                     Quadrupedal walking
                     Whines
                     Wails
                     Panting chuckles
                     Screeches
                     Screams
                     Sits
                     Stand bipedally
                     Social contact w/others
                     Chest beat

. 6-12 months:

                     Ventral ride
                     Dorsal ride
                     Rump-clinging:Infant follows mother with one or
                     both hands clasping hair on rump.
                     Nursing
                     Solo play:Now includes tree climbing.
                     Leave mother
                     Infant restrain
                     Contact with mother
                     Within arm's reach
                     Within 15 feet
                     Social play
                     Eating vegetation:No food prep, just plucking
                     leaves or gnawing.
                     Genital touching:Mother uses genital touching
                     as training method by which infant is urged to
                     crawl, climb, walk, and run (observed only in
                     captives).
                     Walk bipedally
                     Travel quadrupedally
                     Stand bipedally
                     Sit
                     Nurse
                     Push from nipple

  12-24 months:

                     Dorsal ride
                     Ventral carry
                     Travel quadrupedally
                     Rump-clinging
                     Nurse
                     Push from nipple
                     Social play:More vigorous.
                     Solo play:Swinging, twirling and
                     semi-brachiating in trees.
                     Eating vegetation:Now includes stripping
                     leaves.
                     Grooming mother
                     Chest-patting
                     Foliage-whacking
                     Strut-walking
                     Compressed-lip
                     Whines
                     Wails
                     Panting chuckles
                     Screeches
                     Hoot cries
                     Shrieks
                     Temper tantrum
                       screams
                     Howls
                     Pig-grunts
                     Leave mother
                     Mother restrain
                     Contact with mother
                     Within arm's reach
                     Within 15 feet
. 24-36 months:

                     Dorsal ride
                     Travel quadrupedal:Includes canter gait.
                     Ventral carry
                     Nursing
                     Push from nipple
                     Solo play:Tree play increases, becomes more
                     inventive and daring.
                     Social play:Begins including more tree play.
                     Eating vegetation:Includes preparation:
                     wadding, stripping, peeling.
                     Grooming:Includes siblings, peers, and
                     silverback.
                     Whines
                     Wails
                     Panting chuckles
                     Screeches
                     Hoot cries
                     Shrieks
                     Temper tantrum
                       screams
                     Howls
                     Pig-grunts
                     Belch vocalization:Basic disyllabic variant of.
                     Mount
                     Leave mother
                     Mother restrain
                     Contact with mother
                     Within arm's reach
                     Within 15 feet


  Behavior during pregnancy:

                     Proximity
                     Groom infants
                     Staring at infants
                     Contact with mother/infant
                     Distant travel
                     Feeding
                     Mount
                     Copulation
                     Irritability
.Fossey, D.  (1982).  Reproduction among free-living mountain
gorillas. 
                    American Journal of Primatology Supplement,
                    1:97-104.

  Purpose:           To summarize status of and reproduction in
                     free-living gorillas.

  Sampling method:   Ad lib field notes.

  Ethogram key words:Sexual behavior, Field study, Gorilla g.
beringei.
  
  Behaviors:

  Copulation:        Nearly all solicited by females, most commonly
                     dorsoventral.  Male sat upright or leaned
                     forward bipedally and held female around waist. 
                     Female squatted on lap or bent forward on
                     flexed elbows with rump accessible.  Brief
                     adjustment occurred first.  Held position from
                     5 to 20 minutes.  Accompanied by
                     vocalizations, primarily by male.  Female often
                     had compressed lips; male pursed-lips.  

  Copulation vocalization:Panting and soft hoots.

  Solicitation by female:Sometimes included extending hands
                     fleetingly to touch male's body before
                     retreating; often included glance over
                     shoulder.

  Glance over shoulder:To male during solicitation: "come hither
                     look."

  Sexual play

  Pseudosexual mounting

  Interference w/copulation

  Genital touching

. Frisch, D. J.  Columbus Zoo, Gorilla Birth Watch Protocol.

  Purpose:           To recognize changes in behavior of pregnant
                     female, to record onset of labor.

  Sampling method:   Every 15 minutes, note activity of pregnant
                     female.  If unusual behavior occurs between
                     interval, write in time and explain. 

  Ethogram key words:Birthwatch.

  Behaviors:

  Lie:               Being recumbent with little or no weight being
                     borne on hands or feet.

  Sit:               Bearing most weight on buttocks.

  Stand still:       Bearing weight on feet quadrupedally, or
                     bipedally with hands holding on to mesh, glass,
                     trees, etc.

  Locomote:          Walking, running, or climbing.

  ?                  Activity can't be scored.

  Rest near:         Sitting or lying within arm's reach of another
                     for at least 10 seconds.  Initiator is second
                     animal to rest.  The receiver must be
                     relatively immobile but need not be resting.

  Grooming:          Directed touching, licking, or intense visual
                     inspection of another's (GO) or own (GS) skin
                     or pelage.   Scratching excluded.

  Intimidation display:Beating on chest or abdomen, beating on cage
                     surface, rapid exaggerated throwing or shoving
                     of objects.  Score when any two or more of
                     these behaviors occur simultaneously or
                     sequentially.  Score initiator only.

  Rough up:          Slapping, poking, elbowing, punching, hair
                     pulling, pushing, shoving, or wrestling when
                     not in play or sexual context.

  Aggressive bite:   Biting when not in play.

  Crouch:            Initiator lowers ventrum toward or to
                     substrate, or curls fetally on substrate. 
                     Event terminated when normal or resting
                     posture resumed.

  Inspect genitalia: Sniffing, touching, or intense visual
                     inspection of another's genitalia (IGO) or own
                     genitalia (IGS).

  Sex solicit:       Female purses lips and stares
                     at/follows/reaches toward male.  Event 
                     terminated when lips relax for at least 10
                     seconds.

. Copulation:        Dorso-ventral mounting (DVC) or ventro-ventro
                     mounting (VVC) with pelvic thrusting.  Event
                     terminated when either animal breaks body
                     contact.  Note if intromission confirmed (very
                     difficult) or if semen observed afterward on
                     penis or vagina.

  Masturbate:        Rhythmic rubbing of genitalia with any part of
                     own body (e.g., finger or toe) or against
                     inanimate object.

  Yawn:              Per usual definition.  Score each yawn.

  Feces:             Manipulation or mouthing of own or another's
                     feces.  Note "eat" if fecal material actually
                     ingested.

  Regurgitate:       Vomiting into mouth, hand, or onto substrate. 
                     Note if reingested.

  Play behavior:     Poking, tickling, wrestling, chasing in
                     nonaggressive manner.

.Goerke, B., Fleming, L., & Creel, M.  (1987).  Behavioral changes of
a
                                             juvenile gorilla after
                                             a transfer to a more
                                             naturalistic
                                             environment.  Zoo
                                             Biology, 6, 283-295.

  Purpose:           Investigation of one male juvenile lowland
                     gorilla before and after transfer to new
                     environment, particularly play and
                     stress-related behaviors.

  Sampling:          10 minute periods from 10 am to 4 pm daily. 
                     Two indices: play index: number of minutes
                     during which some play occurred in each 10
                     minute observation period.  Frequency: is the
                     number of occurrences in each period of 19
                     selected play activities, divided into social
                     or solitary play.

  Ethogram key words:Post-occupancy evaluation, Environment change,
Influence of, Abnormal/Stereotypic behavior.

  Behaviors:

  19 play activities:

                     Gentle wrestling:Hugging and rolling.

                     Rough wrestling:Standing up, more aggressive
                     behavior.

                     Chasing

                     Running alone

                     Playing with water

                     Slapping

                     Chest pounding

                     Clapping

                     Playing with branches

                     Swinging

                     Jumping

                     Tumbling

                     Spinning

                     Throwing
                     
                     Manipulating objects

                     Moving of the body

                     Solitary sexual play

                     Social sexual play
 
                     Climbing

                     Grass play

                     Dirt play

                     Tree play

  Stress-related behaviors:

                     Coprophagy

                     Regurgitation/reingestion

                     Self-clasping
.Gold, K.  (a)  Noise disturbance study, San Francisco Zoological
               Gardens (currently affiliated with Zoo Atlanta).

  Purpose:To study effects of construction noise on gorilla behavior.

  Ethogram key words:Activity budgets/General behaviors, Noise
disturbance, Effects of.

  Locations:         Trees, rocks, quadrants 1-7, gorilla world, out
                     of sight.

  Behaviors:

  Environmental exploration: Investigation such as touching,
                     sniffing, close visual inspection of parts of
                     the exhibit, habitat, or cage other than
                     gorillas, loose objects, or toys.  Does not
                     include aimless oral or manual activities.

  Eat:               Chewing/ingestion of food/browse.

  Self directed behavior:Self groom, self clasp, self suck,
                     scratching, rubbing, etc.

  Display behavior:  Behaviors used by an animal to draw attention
                     to oneself, oftentimes communicating a specific
                     behavioral state in the displayer.  Includes:
                     
                     Chest beating
                     Ground or wall slapping
                     Drumming
                     Throwing objects
                     Foot stomping
                     Kicking
                     Clapping
                     Branch whipping
                     Strut walking
                     Pursed-lip tense face
                     Other

  Vocalization:      Scream, grunt, bark, etc.

  Solitary behavior: Active physical motions such as in play but
                     without a partner, including rolling, sliding,
                     swinging, spinning, etc.

  Contact:           The focal animal is in direct physical contact
                     with another group member.

  Social play:       The focal animal is engaging in play behavior
                     with another group member (non-aggressive) such
                     as chasing, wrestling, mouthing, etc.

  Aggression:        Chase, hit, grab in aggressive manner
                     (different than social play).

  Sexual behavior:   Oral-genital contact, genital-genital contact, 
                     digital-genital contact, masturbation.

  Locomotion:        Continuous horizontal or vertical movement of
                     at least 1 body length during the interval.

  Rest:              In an inactive state for a minimum of 10
                     seconds.

  Miscellaneous behavior:Digging up enclosure, coprophagy,
                     regurgitation, regurgitation and reingestion.
. Gold, K. (b) Gorilla development study.  Nationwide, 10/82-10/86.

  Purpose:           To investigate infant development in gorillas.

  Sampling:          One/zero sampling during 15 second intervals. 
                     Trial=twelve 15 second intervals with a two
                     minute pause (five minutes total). 
                     Session=Twelve trials (60 minutes).  Focal
                     animal sampling.

  Ethogram key words:Development.

  Behaviors:

  Approach:*         Focal animal moves toward another (or another
                     toward focal) to within proximity.

  Withdraw:*         One animal retreats from another in a
                     non-passive fashion.  Must be the result of an
                     approach.

  Leave by other:**  An animal leaves proximity of another, but not
                     as an result of an approach.

  Touch:*            Physical contact with non-focal animal other
                     than in categories below.

  Lift/hold:**       Weight support contact.

  Carry:**           Infant is transported by another animal.

  Ventro-ventral contact:**Infant's ventrum in contact with other's.

  Dorso-ventral contact:**Infant's ventrum in contact with other's
                     back, shoulders, or back of neck.

  Proximate:**       Within arm's reach of another (arm length of
                     larger animal).

  Near:**            Out of arm's reach of another, but within
                     fifteen feet.

  Social play:*      Physical play with a partner including "rough
                     and tumble" play such as tickling, wrestling,
                     chasing.  Score TOUCH in addition if it occurs.

  Genital inspect*:  Close observation, touching, sniffing or
                     manipulation of genital area of self or other.

  Sexual behavior*:  Mounting, pelvic thrusting, mock copulation,
                     with a partner or inanimate object.

  Nipple contact:**  Infant's mouth on nipple, with or without
                     actual suckling visible.

  Environmental exploration: Investigation such as touching,
                     sniffing,  close visual inspection of parts of
                     exhibit, habitat or cage other than gorillas or
                     loose objects, toys.  Does not include aimless
                     oral or manual activities.

  Object contact:    Contact with toys or any loose object.

  Mouthing:          Using tongue, lips, teeth to explore
                     environment, objects or animals.  May include
                     biting, chewing, food or non-food.  May be
                     during a play bout.

  Solitary play:     Active physical motions such as in play, but
                     without a partner.  Includes rolling, sliding,
                     swinging, spinning, etc.

  Self-directed behavior:Self clasp, self groom, self suck,
                     scratching, rubbing, etc.

  Play face:         Open mouthed, slack lipped "smile" exposing
                     teeth.  May be accompanied by head shake,
                     patting, mouthing or play biting.

  Locomotion:        Continuous horizontal or vertical movement of
                     at least one body length during trial.

  Rest:              In an inactive state for five seconds or more.

  Vocalization:      Belch
                     Cough/grunt
                     Growl
                     Hoot
                     Bark
                     Scream
                     Whine/whinny
                     Pant

  Display:           Chest beat
                     Throwing
                     Slapping/tearing
                     Whipping
                     Thumping/Slamming
                     Strut Walk
                     Threat face

  Out of sight:      Observer is unable to see focal animal.

  Contact with human:Any contact with a person.  Must be initiated
                     by focal animal in order to be scored.

  * note initiator/recipient
  ** note non-focal animal
.Gould, J. E.  University of West Florida and The Zoo, Gulf Breeze,
               Florida.

  Purpose:           Collection of behavior, posture/locomotion, and
                     location data, primarily to document
                     socialization and group formation involving a
                     previously isolated adult male.

  Sampling:          Focal instantaneous time samples, at one minute
                     intervals when observing two gorillas
                     (male-female dyad) and 30 second intervals
                     when observing one gorilla.  These data are
                     supplemented with audiotaped notes, written ad
                     lib notes, and still and video photography.

  Ethogram key words:Activity budgets/General behaviors, Social
behavior, Sexual behavior, Display/Agonistic behavior,
Introductions/Socialization, Abnormal/Stereotypic behavior,
Vocalizations.

  Behaviors:         

  Positional Behaviors:

  Posture:

   Lie:
   
    Lying down:      Specify dorsal, ventral, lateral.

    Leaning:         Legs under body and body less than 45 degrees
                     from horizontal--specify ventral or lateral and
                     on which arm or arms.

   Sit:              

    Sitting:         Resting on buttocks--specify any support or
                     suspension with hands.

    Squatting:       Knees bent, buttocks above heels--specify any
                     support or suspension with hands.

    Standing, quadrupedal
                     /pronograde

    Standing, tripedal

    Standing,
                     bipedal/upright

    Crouching:       Quad stand with limbs flexed/bent.

  Locomotion:        

   Walk:             

    Walking, quadrupedal
                     /pronograde: (knuckles?)

    Walking, tripedal

    Walking,
                     bipedal/upright

   Run:              

    Running, quadrupedal
                     /pronograde

    Running, tripedal

    Running,
                     bipedal/upright

   Other:

    Climbing:        Specify up, down, left or right.
    Hanging by hand(s)

    Swinging by hand(s)

    Brachiating:     Progressing by swinging from one hold to
                     another by the arms.
    Jumping

    Spinning

    Sliding

    Rolling:         Sideways.

    Somersaulting:   Head over heels.

  Eating, drinking, and eliminating activity:

   Eat:              

    Eating:          Specify type of food.

    Gathering:       Gathering food in heaps.

    Foraging:        Wandering in search of food.

    Sharing food:    With another gorilla.

    Symbolically eating

   Drink:            

    Drinking water

    Drinking milk

    Nursing
   
   Eliminate:        

    Defecating
 
    Attending to feces

    Handling feces

    Passing gas

    Urinating

    Attending to urination

    Handling urine

  Object-oriented activity (specify if play):

   Look (specify object/food):

    Glancing:        Briefly observing sideways out of corner of
                     eyes.

    Looking:         Observing directly.

    Watching/staring:  Prolonged and attentive observing; curiosity?

   Manipulate (specify object/food and body part used): 

    Smelling

    Mouthing

    Reaching

    Touching
    Fiddling

    Grasping/grabbing

    Clasping/hugging/
                     cuddling/embracing:In arms.

    Releasing

    Patting

    Swiping

    Slapping

    Hitting

    Kicking

    Barging

    Pushing

    Pulling
 
    Throwing

    Swishing/splashing: Water.

    Holding:         Specify hands, feet, mouth, back, etc.

    Carrying:        Specify hands, feet, mouth, back, etc.

   Other:            

    Approaching      

    Moving away

    Retreating:      Withdrawing from apparent danger (e.g., snake).

    Avoiding/evading

    Fleeing/escaping

    Grimacing:       e.g., at water spray.

    Swatting insects

    Killing insects

    Catching insects

    Nest building

    Tool using:      Specify tool and object acted on.

    Tool manufacturing:Specify tool and method.

  Self-oriented activity (specify if play):

  Manipulate:        

   Non-genital self-manipulating:

    Autogrooming:    Picking through own hair with fingers and/or
                     lips.

    Rubbing

    Scratching

    Picking:         Specify nose, eye, etc.
    Finger tapping

    Hand clapping

    Chin slapping:   See chest-beating sequence below.

    Hair pulling:    Stereotypic/abnormal?

   Genital self-manipulating:

    Masturbating

    Bottom bouncing:   By female.

    Self-inspecting:   Visually.

    Finger licking:  May be associated with menstruation.

    Finger smelling: May be associated with estrus.

Stereotypic/Abnormal activity (may not be abnormal unless behavior
is stereotypic; could include preceding behaviors listed under
"manipulate"):

   Rocking           

   Swaying:          Side to side.

   Shuffling

   Pacing

   Self-biting

   Self-grasping

   Turning back

   Sitting in corner:  As though in bed.

   Jawing:           Opening and closing mouth.

   Grinding teeth

   Regurgitating and
    reingesting

   Coprophagy:       Eating dung.

   Drinking urine

  Other:             
   
   Sleeping:         Eyes closed, no movement for 3 minutes.

   Resting:          Specify if eyes are closed--see sleeping.

   Yawning

   Looking:          At self; specify body part if possible.

   Self recognition: In mirror or other reflector.

   Sitting or standing
    in water

   Running through water

   Splashing in water

   Chest-beating sequence:Release of accumulated tension associated
                     with excitement as in sexual excitement, play,
                     mild irritation, impatience, frustration,
                     lonesomeness, novelty, or when in conflict
                     between aggression and flight; this is the
                     apparent primary function and cause; for
                     additional functions as well as descriptions
                     and components to be specified--see below under
                     "inter-individual oriented activity" both
                     "interact" and "aggress-defend."

  Inter-individual oriented activity (specify if play):

   Look:

    Glancing:        Briefly observing sideways out of corner of
                     eyes.

    Looking:         Observing directly.

    Watching:        Prolonged and attentive observing.

    Staring:         Listed below under "reproductive" and
                     "aggressive acts".

  Interact (nonagonistic behavior, includes play; note that several
  of these behaviors also occur in agonistic interactions--see below
  under "aggress-defend"):

    Non-contact:

                     In vicinity:1-5 meters.

                     In proximity:Within arm's length; approximately
                     1 meter.

                     Approaching:Moving towards.

                     Displacing:Taking place of another.

                     Lunging:Incipient charge: abrupt advance of 0-3
                     steps.

                     Charging:Running is lateral, not forward, if
                     there is conflict between charge and retreat.

                     Chasing:Pursuing rapidly.

                     Throwing material:Specify material.

                     Standing off:Rigid/strut quad stance, facing
                     each other; often associated with tight-lipped
                     face--see below under "other," can be sexual
                     initiation by female toward male or vice versa.
                     Moving away

                     Retreating:Withdrawing from what is dangerous
                     or disagreeable.

                     Avoiding/evading

                     Fleeing/escaping

                     Chest-beating
                        sequence:See above.

                     Reaching/inviting:Arm extended, hand open, palm
                     up; initiates play; see also under
                     "reproductive".

                     Arm-over response:Palm down.

                     Smelling

                     Imitating

                     Teasing

                     Playface/pleasure
                       face:Relaxed, open mouthed.

                     Play biting

                     Play walking

                     Play running

                     Play chasing

                     Play fleeing

                     Bouncy gait

    Contact:

                     Touching:Non-genital.

                     Hand holding

                     Grasping/grabbing

                     Clasping/hugging/
                      cuddling/embracing: In arms.

                     Releasing

                     Giving

                     Taking

                     Leaning on

                     Grooming:Picking through hair of another with
                     fingers  and/or lips.

                     Tagging

                     Patting

                     Swiping

                     Slapping

                     Hitting

                     Kicking

                     Pummelling

                     Barging

                     Pushing

                     Pulling

                     Wrestling/grappling

                     Mouthing:Taking an arm or shoulder between
                     teeth and appearing to bite, but applying no
                     pressure; can be intimidation by dominant male
                     and thus agonistic.

    Reproductive:

                     Charging parallel:By male, sometimes preceded
                     by chest-beating display, usually followed by
                     standing off.

                     Standing off:See above under "non-contact".

                     Staring:Can be sexual initiation by female
                     toward male, often associated with standing off
                     and head shaking, may be tight-lipped.

                     Head shaking:See below under "aggress-defend:
                     submissive displays".

                     Inviting:Extended arm with hand open and palm
                     up by female while staring at male; another
                     variation is drumming on hindquarter or ground
                     by female while staring at male.

                     Offering/presenting:Prone position: female
                     crouches, backs up toward male to display her
                     genitalia, and looks back at him; supine
                     position: female lays on back and makes
                     rhythmic pelvic movements, also often inviting
                     as above.
                     Inspecting genitalia

                     Manipulating gen.

                     Oral-genital contact
 
                     Mounting:Specify dorso-ventral or
                     ventro-ventral.

                     Pelvic thrusting

                     Paternal/maternal
                       caring:e.g., nursing.

                     Disciplining

  Aggress-Defend (agonistic behavior, aggressive or defensive
interaction--see also  behaviors listed above under "interact"):

    Aggressive acts:

    Non-contact:
    
                     Staring:Furrowed brow?

                     Staring openmouthed
                      /threat face: Specify if teeth showing.

                     Staring bared-
                      teeth scream face/
                      threat face
                     
                     Head jerking or
                      snapping:Toward antagonist; specify if audible
                     snap of jaws.
                     
                     Forward lunging:Incipient charge: abrupt
                     advance of 0-3 steps.
                     
                     Bluff charging:If conflict between charge and
                     retreat; then lateral running.

                     Throwing non-fecal
                        material

                     Throwing feces

    Physical contact:  

                     Hitting
                     Barging
                     Grappling
                     Biting
                     Etc.:See "interact: contact" behaviors above.

   Threat Displays (Domination):

    Chest-beating
                     sequence:Intimidation to repel intruders; for
                     functions other than aggress-defend, see above
                     under "interact" and "self-oriented activity". 
                     Consists of rhythmic beating, with alternating
                     cupped hands, of the chest, but also abdomen
                     and thighs, as well as cheek drumming with
                     mouth open, chin drumming with backs of 
                     fingers, and rhythmic beating of ground, floor,
                     logs, etc.; chest-beating is only one of 9
                     actions which do not necessarily all occur
                     together, or in the following order; full
                     sequence exhibited only by silverbacked males
                     and infrequent--specify components present:

                       Hooting
                       Symbolically feeding
                       Rising bipedally
                       Throwing vegetation
                       Chest-beating
                       Leg kicking
                       Running sideways bipedally then quadrupedally
                       Slapping and tearing vegetation
                       Ground thumping (or beating/slapping other
                     inanimate object, e.g., wall)

    Strut standing:  Very stiff quad posture, elbows bent outward,
                     shoulders hunched, sometimes head diverted to
                     side, brief sideways glances at opponent;
                     commonly with tight-lipped/lip-in/tense-mouth
                     face; may occur at end of chest-beating
                     display.

    Strut walking:   As above, with short-stepped quadrupedal
                     walking.

    Strut running:   May be followed by striking a barrier or
                     another animal with shoulder or forearm.

   Submissive displays (Submission):

    Turning head away: Also used to indicate aggression is not
                     intended.

    Head shaking:    Also used to appease when unsure and during
                     sexual initiation by female while staring at
                     male; relatively rare; specify if lower jaw
                     relaxed.

    Cowering:        Crouching on abdomen; head lowered, arms and
                     legs tucked under; protects body.

   Other:

    Pursed-mouth/pout/
                     light-distress face:Watch out, I'm ready to get
                     mad; or male during copulation.

    Tight-lipped/lip-in/
                     tense-mouth face:Usually with periodic quarter
                     turns of head and sideways glancing; situations
                     of uncertainty, tension, courtship; or female
                     during copulation.
    Grimacing
 
    Fear grinning

    Vocalizing:      Specify, see vocalizations below.

    Piloerecting:    Erecting body hair, especially on back and
                     head.

    Marking:         e.g., by urinating.

  Observer-oriented activity (specify if play):

   Look:

    Glancing:        Briefly observing sideways out of corners of
                     eyes.

    Looking:         Observing directly.

    Watching/staring:  Prolonged and attentive observing; curiosity?

  Interact (non-agonistic behavior, includes play; see
"inter-individual oriented activity" above for additional behaviors
and descriptions):

    Listening

    Pursed-mouth/pout/
                     light-distress face

    Tight-lipped/lip-in/
                     tense-mouth face

    Chest-beating sequence

    Vocalizing:      Specify, see "vocalizations" below.

    Exchanging/giving

    Imitating

    Signing:         Specify: e.g., chest tap, head tap, hand clap.

  Aggress-Defend (see "inter-individual oriented activity" above for
listing of specific    behaviors):

    Aggressive Acts

    Threat displays
    (Dominance)

    Submissive displays
    (Submission)

  Vocalizations:

   Aggressive Calls:

    Roar/threat bark:  Simple, mono-syllabic, sudden loud outburst
                     of low-pitched harsh sound.  Strong aggression
                     of silverback male to predator (including
                     human) or other group.  Always followed by some
                     type of aggressive behavior.

    Growl:           Deep guttural prolonged noise resembling growl
                     of dog and not loud.  Mild aggression in
                     stationary group associated with "annoyance."

    Pant series:     Rapid sequence of forced expulsions of air, of
                     very low frequency, seems whispered, resembles
                     pig grunt (see below).  Mild threat within
                     group.

   Mild alarm calls:

    Question bark:   Three notes, low-high-low, as in "who are you?" 
                     Very mild alarm or curiosity.

    Hiccup bark:     Resembling di-syllabic spurt of noise like a
                     hiccup.  Very mild alarm of curiosity.

   Fear and Alarm Calls:
    
    Silence:         Sudden cessation of all sound emission, vocal
                     and mechanical.  Most extreme form of fear,
                     e.g., to sounds of approaching poachers.

    Screams:         Shrill and prolonged emission of extremely loud
                     sound.  Aggressive disputes within group;
                     copulating female.

    Wraagh/alarm bark: Explosive, monosyllabic outburst of loud
                     sound, abrupt beginning and end.  Sudden
                     alarming situation; loud noise; very close or
                     sudden contact; not accompanied by aggression.

   Distress calls:

    Cries:           Wails, sobs, screeches and hooting types of
                     distress calls; can build to screams or shrieks
                     (temper tantrums).  Infant separated or in
                     difficulty.

    Whines:          Very soft, prolonged plaintive sound, like that
                     of puppy.  Danger of injury or abandonment; not
                     limited to infants.

   Group coordination vocalizations:

    Pig grunts:      Short, rough, guttural grunts (av. 9-10),
                     resembling grunting of pigs feeding at a sty. 
                     Mild aggression in moving group, e.g., right of
                     way dispute during feeding; silverback when
                     pursuing female before copulation or when
                     guiding group movements; disciplinary 
                     enforcement.

    Hoot bark:       Like bark of dog but more "who-who" than
                     "woof-woof" or "wah-wah" sound; like hiccup and
                     question barks but two, not one or three
                     syllables.  Alerting to mild alarm; group
                     movement initiation.

    Belch vocalization:Deep, prolonged rumble, like soft belch; most
                     commonly long, di-syllabic throat-clearing
                     (ummm-ummm) sound, with second note lower than
                     first; at times more complex, like croons,
                     purrs, hums, moans, wails, or howls.  Feeding;
                     group contentment; localization; useful in
                     allaying gorilla apprehension among human
                     observers.

.  Intergroup communication:

    Hoot series:     Given with or without a terminating chest-beat
                     or other elements of that sequence.  Prolonged
                     series (av. 10-15) of "hoo hoo hoos",
                     low-pitched, quiet at start of series, building
                     up.  Intergroup encounters with aggressive
                     component; intergroup communication/vocal
                     probing/social space echolocation.
   
   Miscellaneous vocalization:

    Chuckles:        Raspy expirations verging between pants and
                     grunts.  Social play, tickling.

    Copulatory pants:  Rapid series of soft, short duration hoots,
                     becoming more prolonged and merging into long
                     howl-like note.  Silverback copulating.  Also
                     dove-coos by both sexes.

    Whinnies:        Resembles raspy neighing of horse.  May be
                     anomalous; ailing animal.
.Harcourt, A. H.  (1979a).  Social relationships among adult female
                         mountain gorillas.  Animal Behaviour,
                         27:251-264.

  Purpose:           To examine relationships among adult female
                     mountain gorillas.

  Sampling:          Ad lib sampling, using behavioral checksheet,
                     including intitiator/recipient, bout lengths. 

  Ethogram key words:Social behavior, Field study, Gorilla g.
beringei.
  
  Behaviors:         (Also see Harcourt, A. H. (1977).  Social
                     relationships of wild mountain gorilla.  Ph.D.
                     thesis, University of Cambridge.)

  Proximity:         Within 2m or 5m of partner.

  Touching:          More than 5 sec.

  Body contact:      Excludes grooming.

  Grooming:          More than 5 sec.

  Agonism:

                     Avoid:Actions ranging from simply leaning away
                     from an approaching animal to cringing under an
                     attack or getting up and leaving.

                     Pig-grunts:Antagonistic, coughing-type
                     vocalization.

                     Chest beating

                     Strut runs

.Harcourt, A. H.  (1979b).  Social relationships among adult male and
                         female mountain gorillas in the wild. 
                         Animal Behaviour, 27:325-342.

  Purpose:           To examine relationships among adult male and
                     female mountain gorillas.

  Sampling:          Continuous ad lib sampling, using behavioral
                     checksheet, including intitiator/recipient,
                     bout lengths; plus activity and location scans
                     at 15 minute intervals.

  Ethogram key words:Social behavior, Field study, Gorilla g.
beringei.
  
  Behaviors:         (Also see Harcourt, A. H. (1977).  Social
                     relationships of wild mountain gorilla.  Ph.D.
                     thesis, University of Cambridge.)

  Proximity:         2m and 5m.

  Grooming:          Only those lasting > 5 sec.

  Agonism:

                     Approach-retreat:Non-aggressive approach
                     followed by retreat.

                     Approach

                     Retreat

                     Avoid:Movement away within 1 to 2 sec following
                     the other's approach, but not to beyond 2 m

                     Supplanted:Movement away within 1 to 2 sec
                     following the other's approach, to beyond 2 m.

                     Threat display:Includes chest-beating,
                     ground-thumping, foliage-slapping, strutting
                     and running.

                     Hooting

                     Attacks:Hits, kicks, and bites.

                     Pig grunt:Sharp grunt.

.Harcourt, A. H. & Stewart, K. J.  (1981).  Gorilla male
                                             relationships: Can
                                             differences during
                                             immaturity lead to
                                             contrasting
                                             reproductive tactics in
                                             adulthood?  Animal
                                             Behaviour, 29:206-210.

  Purpose:           To examine relationships among adolescent males
                     and group leaders and tendency to remain in
                     natal group.

  Sampling:          Ad lib sampling, using behavioral checksheet,
                     including intitiator/recipient, bout lengths. 

  Ethogram key words:Social behavior, Field study, Gorilla g.
beringei.
  
  Behaviors:         (Also see Harcourt, A. H. (1977).  Social
                     relationships of wild mountain gorilla.  Ph.D.
                     thesis, University of Cambridge.)
                     (Also see Stewart, K. J.  (1981).  Social
                     development of wild mountain gorillas.  Ph.D.
                     diss., University of Cambridge.)

  Proximity          
  Contact            
  Grooming           

.Hoff, M. P., Nadler, R. D., & Maple, T. L.  (1981a).  Development of
                                                       infant
                                                       independence
                                                       in a captive
                                                       group of
                                                       lowland
                                                       gorillas. 
                                                       Developmental
                                                       Psychobiol-
                                                       ogy, 14,
                                                       251-265.

  Purpose:           To supplement the data available on infant
                     development of gorillas.

  Sampling:          One/zero sampling, one minute intervals in 1st
                     3 and last 5 months; focal animal continuous
                     sequential activity record for remainder of the
                     18 month study using 30 minute samples.

  Ethogram key words:Development.

  Behaviors:

  Ventro-ventral:    Animals' ventral surfaces in contact.

  Dorso-ventral:     One animal's ventrum pressed against another
                     animal's dorsum.  Herein used exclusively in
                     "back ride" position (infant lying on mother's
                     back).

  Ventral-dorso:     As above.  Herein used exclusively as infant's
                     dorsum against its mother's ventrum.

  Other contact:     Any form of contact excluding ventro-ventral,
                     dorso-ventral and ventro-dorsal.

  Proximate:         Animals not in contact, but within 3 ft (1 m)
                     (arm's length) of one another.

  Distant:           Animals between 3 and 15 ft (1 and 5 m) apart. 

  Far distant:       Animals beyond 15 feet (5 m) apart.

  Leave:             Moving from a position of contact to proximate,
                     distant or far distant.

  Return:            Reestablishing contact with another animal. 
                     Scored for the animal reestablishing contact,
                     regardless of which animal left.

  Social approach:   Animal moving from distant or far distant to
                     proximate (contact not established).

  Follow:            Moving behind and in the same direction as
                     another animal.

  Restrain:          Prevent another animal from moving away.

  Struggle:          Animal attempting to leave another while being
                     restrained.

  Retrieval:         Move another animal into closer spatial
                     proximity as defined by the above contact 
                     variables.

  Nipple contact:    Animal's mouth placed on another's nipple with
                     or without suckling.

  Wean:              Remove an infant from the nipple.

  Remove:            Move another animal into a position of less
                     close spatial proximity as defined by the above
                     contact variables.

  Social examination:Inspect another animal with fingers, feet,
                     and/or mouth (includes stare).

  Object examination:Inspection and/or manipulation of articles in
                     the environment (does not include solitary play
                     using objects).

  Protect:           Saving an animal from possible harm (by
                     retrieval, crouching over the animal, holding
                     arm up, etc.).
.Hoff, M. P., Nadler, R. D., & Maple, T. L.  (1981b).  Development of
                                                       infant play
                                                       in a captive
                                                       group of
                                                       lowland
                                                       gorillas
                                                       (Gorilla
                                                       gorilla
                                                       gorilla). 
                                                       American
                                                       Journal of
                                                       Primatology,
                                                       1, 65-72.

  Purpose:           To supplement the data available on infant
                     play, and particularly of development of infant
                     play in gorillas.

  Sampling:          One/zero sampling, one minute intervals in 1st
                     3 and last 5 months; focal animal continuous
                     sequential activity record for remainder of the
                     18 month study using 30 minute samples.

  Ethogram key words:Development.
  
  Behaviors:

  Infant solitary play:Initially consisted of ground slapping...at
                     7th month consisted of gymnastic play.

  Mother-infant play:Typically involved gently nudging with finger,
                     arm, or leg by the mother, and wrestling with
                     and mouthing of the mother by the infant.  Play
                     was often exhibited in a discontinuous,
                     reciprocal fashion.

  Infant-infant play:

                     Active social play:"Rough and tumble" play. 
                     Typically consisted of chasing, lunging,
                     tackling, vigorous wrestling, falling on the
                     other, and vigorous mock biting.

                     Moderate social play:Typically involved lying
                     on the play partner, light bouncing on the
                     partner's stomach, or mild pulling.


.Lockard, J.  S.  Ethogram for longitudinal study of Seattle's
                    Woodland Park gorillas.  University of
                    Washington.

  Purpose:           To develop a longitudinal database on captive
                     gorillas.

  Sampling:          15 minute continuous focal-animal samples,
                     divided into 30, 30-second intervals, of all
                     behaviors; with location scans at 5 minute
                     intervals.

  Ethogram key words:Activity budgets/General behaviors, Social
behavior, Sexual behavior, Development, Maternal behavior,
Display/Agonistic behavior, Vocalizations.

  Behaviors:         (Only an abbreviated ethogram is presented
                     here.  A manuscript describing the full
                     ethogram, and detailing the data collection
                     process, is presented in Appendix 1.)

  NonSocial Behaviors:

  Locomotive Behaviors:
                     
                     Quadrupedal:

      Stand:         On all four extremities.

      Walk:          Forward progression or backward motion.

      Lateral walk:  Forward progression with sideways deviation
                     (species characteristic).

      Run:           (Conventional definition.)

    Tripedal stand/walk/
      run:           On three extremities.

    Bipedal stand/walk/
      run:           On two lower extremities.

    Sit:             Buttocks in contact with ground or supported by
                     heels with the back vertical.

    Handstand:       (Conventional definition.)

    Lie:             Reclines on substrate.

    Circle:          Spin, roll, somersault in self play
                     (conventional definitions).

    Climb:           Movement up/over an object.

    Jump:            Vertical spring from substrate.

    Fall:            Tumbles abruptly from one position to another.

    Hang:            Suspended by one or both arms from elevated
                     surface.

    Scoot:           Swings legs between hand placement while in a
                     sitting posture.

    Slide under:     Squeezes beneath object in reclined position.

  Self-directed Behaviors (while stationary):

    Urinate/defecate:(Conventional definitions.)

    Collect/discard stool:Manually gathers feces eats/or drops.

    Cradle foot:     While reclined, holds lower extremity in arm.

    Self maintenance:Manual grooming of face and body.

    Rest:            Reclined with eyes closed.

    Pat:             Hit self with hand.
    Regurgitate:     Manually collects vomit, eats/or discards.

    Shake:           Repeatedly moves hand/or head in self play.

    Adult self nurse:Mouths own nipple.

  Food and/Environment Directed Behaviors:

    Consume:         Eat/drink.

    Forage:          Search for, uproot, tear off and/or pick up
                     edible items.

    Chase birds:     Rapidly following fleeing individual(s).

    Pull/rock object:Manipulates object.

    Pry:             Attempts to open door/window with hand or
                     object.

    Thread:          Attempts to manually insert object through
                     opening.

    Drape:           Covers body part with bough or object (e.g.,
                     burlap).

    Nest build:      Arranges objects (straw, branches, burlap) in
                     rim around and/or under body.

    Hold object(s):  Supports objects in mouth or extremity(ies).

    Lick/sniff:      Tongue touch or smells object.

    Shred object:    Tears apart or peels object with hands or
                     mouth.

    Rub:             Touches object with back and forth motion.

    Drop object(s):  Releases object from mouth or extremity(ies).
 
    Tool dig:        Uses object to disturb substrate.

    Lift/sift:       Manually scoops soil and discards.

    Brush substrate: Smooth and/or clear area with hand.

    Look at/up:      Eyes fixed on (environmental) object.

  Noncontact Social Behaviors:

  Facial displays:

    Yawn:            Tips head back, exposing mouth cavity fully or
                     partially while inhaling.

    Attention:       Active, direct or over-the-shoulder observation
                     of conspecifics/humans in area.

    Relaxed face:    Eyes neutral and mouth closed, surveys
                     environment and visible other(s) without
                     obvious attention.

    Estrus gaze:     Hard, fixed gaze by female conspecific toward
                     male conspecific.

    Stare face:      Head tipped slightly downward, eyes hard and
                     fixed.  Lips pursed or curled back, may scream
                     or roar.

    Uneasy face:     Refrains from direct stare at other.  May have
                     mouth tightly compressed and or head turned to
                     side.

    Pout face:       Lower lip pursed and protruding slightly. 
                     Pulled down lip corners and hollow cheeks may
                     be seen.

    Fear grimace:    Head tilted slightly back, eyes darting back
                     and forth.  Lips curled back at mouth corners,
                     exposing gums and teeth.

    Sneer:           Upper lip drawn back.

    Head shake:      Moves head back and forth.

    Play/laughter face:Mouth open and corners rounded.  Exposed
                     teeth and gums, and laughing motions without
                     vocalizations may be seen.

  Vocalizations:

    Roar:            Sudden, loud outburst of low-pitch sounds from
                     open mouth.

    Growl:           Deep, guttural prolonged sound.

    Pant:            Repeated, rapid exhalations.
 
    Bark:            Dog-like sound.

    Scream:          Shrill, loud, prolonged sound.

    Wraagh:          Abrupt, monosyllabic explosion of loud sound.

    Pig grunt:       Series of short, rough guttural pig-like
                     sounds.

    Belch:           Soft, deep prolonged rumble.

    Hoot:            Series of distinct hoo-hoo sounds.

    Chuckle:         Series of raspy, successive expirations.

    Sexual grunt:    By male when copulation is averted/terminated
                     before fruition.

    Raspberries:     Lips extended, rapid exhalation.

    Staccato grunt:  Similar to pig grunt.  Grunts emitted on
                     inhalation as well as exhalation.

    Hoot-whine:      High volume hoot with level dropping
                     considerably to a narrow band of prolonged
                     sound.

  Postural displays:

    Strut displays:  Stiff quadrupedal stance, arms bent outward at
                     elbow (species characteristic).  Frontal
                     oblique, or parallel variants.

    Strut display w/motion:As defined above, while walking alone or
                     in parallel with other.

    Running display: Bipedal or tripedal run with outswept arm (hand
                     holding object or empty).

    Reach for:       Extends extremity in direction of other.

    Throw:           Manually tosses object.

    Catch:           While stationary, grabs thrown object.

    Arm swing:       Sweeps arm out towards other.

    Chest/body pound:Rapidly beats body with open, slightly cupped
                     hands.

    Kick at:         Rapidly extends hindlimb out and back in
                     direction of other.

    Hit surface:     Strikes substrate/structure with hand or
                     object.

    Social forage:   Coordinated foraging with other(s).

    Social sniff:    Explores body part, excreta or secretion of
                     other.

    Wave:            Flaps hand or arm in air.

    Hand clap:       Brings palms together rapidly.

    Head jerk:       Moves head quickly in direction of other.

    Lunge:           Rapid, short quadrupedal jump toward other.

    Self protect:    At approach of other, cowers, flinches or
                     blocks body with arm.

    Approaches:      Runs or walks to stationary other.

    Leaves:          Runs or walks from stationary other.

    Chases:          Behind other, both running.

    Conflict intervention:Supports one of two (or more) disputing
                     others.

    Leads/follows:   In front of other, both walking.

    Supplant:        Concedes location to approaching other.

    Proximity:       Other in nearby location (within 30-60 cm).

  Contact Social Behaviors:

  Agonistic/play interactions:

    Strike:          Use of extremity in brief, sharp contact with
                     other; e.g., hit, poke, kick.

    Grab:            Manually seize other suddenly.

    Wrestle:         Grabs and hits other.  May roll on ground, each
                     individual grappling, mouthing and slapping
                     other.

    Mockbite:        Teeth in contact with other without
                     penetration.

    Bite:            Seizes other with teeth.

    Push/pull:       Shoves away/draws other closer with extremity.

    Steal:           Grabs object from possession of other.

    Body press:      Uses chest to restrain crouched other.

  Sexual interactions:

    Present:         Remains stationary with ano-genital area 
                     towards other.

    Probe:           Explores genital area of other with hand or
                     mouth.

    Probe avert:     Activity that foils successful probe; sitting,
                     distraction, cowering, threatening, or exiting.

    Mount:           Assumes position for copulation dorsal-ventral,
                     ventral-ventral, or variation.  Male may use
                     hind limb in restraint of female.

    Mount avert:     Activity that foils successful mount; sitting,
                     distraction, cowering, threatening, or exiting.

    Copulation:      (Conventional definition) with multiple
                     thrusts; may see ejaculatory pause.

  Adult/young interactions:

    Nurse:           Suckles nipple(s) of female.

    Nurse prevent:   Activity that prevents young from nursing;
                     blocking nipple, nipping at, and/or pushing
                     away.
    Hold infant:     Support young against body with arm(s) while
                     stationary.

    Hold/carry solicit:Young holds arms above head in proximity to
                     older other.

    Carry infant:    Supports young against body with hand/arm while
                     moving; variants (e.g., dorsal carry, leg
                     carry).

    Pick up:         Manually lifts young.

    Release:         Relinquishes hold on young.

    Shift:           Moves young to another position on body.

    Retrieve:        Pulls young back.

    Shake:           Abrupt manual movement of young.

    Kidnap:          Grabs young from other.

    Infant cling:    Young self-supported by holding onto other.

    Drag infant:     Manually pull infant while moving along
                     substrate.

    Swing:           Rhythmically moves suspended young by arms.

  Other contact affiliative interactions:
 
    Allogroom:       Uses mouth/or hand to inspect/pick pelage of
                     other.

    Social lick:     Mouth to body part, excreta or secretion of
                     other.

    Caress:          Gently rubs mouth or hand over body part of
                     other.

    Repose together: Sit or recline in contact with other.

    Tickle:          (Conventional definition.)

    Kiss:            Mouth briefly touches facial area of other.

    Hug:             Embraces other with arm.

    Social climb:    Ascends/or descends other.

    Fall off:        Loses hold on other.

    Jump on/off:     Abrupt ascent/descent of other.

    Solicit:         Begs food or object from other.

    Offer:           Extend food or object to other.
Mannchen, K.  (1980).  A method of analysis of the activity patterns
                         of lowland gorillas (Gorilla gorilla
                         gorilla) at the Miami Metro Zoo.  Animal
                         Keepers' Forum, 465-471.

  Purpose:           To determine if there were differences in
                     behavior of 5 lowland gorillas.

  Sampling:          One/zero Hansen's checklist, 15 minute samples,
                     100 samples per gorilla.

  Ethogram key words:Activity budgets/General behaviors.

  Behaviors:

  Activities:

                     On ground
                     Climbing
                     Elevated
                     Walking
                     Running
                     Sitting
                     Standing on all fours
                     Kneeling on elbows
                     Standing bipedally
                     Lying down
                     Yawning
                     Sleeping
                     Manipulating with hands
                     Manipulating with mouth
                     Wearing something on head
                     Drinking
                     Urinating
                     Eating
                     Bowel movement
                     Nesting
                     Grooming self

  Social interactions:

                     Grooming another
                     Contact passive
                     Contact active
                     Playing
                     Transport
                     Nursing

  Sexual responses:

                     Masturbation
                     Presenting
                     Copulation
                     Ejaculation
                     Initiating sexual encounter

. Emotionally aggressive and submissive behaviors:

                     Thigh beating
                     Chest beating
                     Charging display
                     Tight lips
                     Strutting walk
                     Beating on objects
                     Throwing objects
                     Grinding teeth
                     Smashes object
                     Pucker hooting
                     Grumble
                     Staring
                     Chasing
                     Hitting
                     Grappling
                     Mock biting
                     Biting
                     Screaming
                     Coalitions
                     Turning of head
                     Head shaking
                     Cowering
                     Dominant supplanting
                     Subordinate supplanted

  Abnormal behaviors:

                     Stereotyped rocking
                     Coprophagy
                     Regurgitation reingestion
                     Pacing
. Maple, T. L. & Hoff, M.  General Gorilla Ethogram.

  Ethogram key words:Activity budgets/General behaviors, Social
behavior, Sexual behavior, Development, Maternal behavior,
Display/Agonistic behavior.

  Behaviors:

  Contact variables:

                     Ventro-ventral:Common usage.

                     Dorso-ventral:M dorsal, I ventral (i.e.,
                     back-ride position).

                     Ventro-dorsal:Vice-versa above.

                     Other contact:Any contact other than above or
                     "touch".

                     Proximate:Out of contact, within arm's
                     (Mother's) length of other.

                     Distant:Farther than one arm's length from the
                     other, less than 15 feet.

                     Far distant:Beyond 15 feet from other animal.

                     Hold:Animal clinging to another.

                     Touch:Momentary contact with hand (this
                     behavior is scored independently of other
                     contact variables).

  Maternal behaviors:

                     Back ride:Infant riding on her mother's back,
                     "jockey style."

                     Hanging walk:Animals VV, mother walking
                     quadrupedally.

                     Tripedal walk:Mother holding infant with one
                     arm while walking.

                     Crutch walk:Infant in mother's lap, M move
                     using arms as crutches.

                     Protect:Save from possible harm; by retrieve,
                     crouching over infant, holding arm up, etc.

                     Retrieve:Move another animal into closer
                     contact.

                     Restrain:Prevent another from moving away.

                     Wean:Remove infant from nipple.

                     Punish:Mother preventing infant from moving
                     into closer position with her (includes pushing
                     animal into the ground).

                     Extended arm carry:Mother carrying infant away
                     from her body and  off the ground.

                     Extended arm drag:Mother carrying infant away
                     from her body but along the ground.

  Infant behavior:

                     Whimper

                     Scream

                     Nipple contact:Infant's mouth on mother's
                     nipple.

                     Struggle:Infant attempts to leave while mother
                     restrains.

                     Arm out:Infant holding arm out to mother (i.e.,
when mother moves away).

                     Crawl:Animal moving on hands and knees.
  Individual behaviors:

                     Active play

                     Solitary play

                     Auto groom:Pick through own hair with fingers
                     and/or lips.

                     Self manipulation:As manipulate but
                     self-directed.

                     Eat

                     Defecate

                     Urinate

  Postural/movement:

                     Lie

                     Sit

                     Squat

                     Quadrupedal stand

                     Quadrupedal walk

                     Bipedal stand

                     Bipedal walk

                     Bipedal walk w/support:Walking bipedally while
                     holding some supporting substrate.

                     Climb
 
                     Leave:Move from other.

                     Social approach:Move from Distant to Proximate.

                     Return:Move back into contact within 15 seconds
                     of leaving.

                     Follow:Moving after another animal.

                     Chase:Following another animal at a run
                     (usually in play).

                     Displace:Take another's place.

                     Retreat:Move away from another as it approaches
                     and follows.

                     Turn away:Turn bodily away from another as it
                     approaches.

                     Walk by:One animal walks by another, becoming
                     proximate or in contact only for a moment.

                     Crawl:Animal moving on hands and knees.

                     Stand with support:Animal standing bipedally
                     while holding some supporting substrate.

                     Perch:Animal off the ground and not moving
                     (i.e., lie on climbing apparatus).

                     Run

                     Gallop

                     Arm-in-arm:Hip hold.  Animals walking
                     tripedally holding each other around the body
                     shoulder (arms entwined).

  Play behaviors:

                     Initiate social play:Animal responsible for
                     beginning social play bout.

                     Passive social play:Less than whole body
                     involvement, often gentle rocking, mouthing, or
                     wrestling.

                     Active social play:Whole body involvement,
                     including rough wrestling, rolling, chasing.

                     Play hit:Animal standing or sitting and
                     repeatedly hitting another (usually during
                     active social play).

                     Side-by-side play:Parallel play - animals
                     proximate and both engaged as in active social
                     play.

                     Play movement:Attempted play imitation
                     (non-reciprocated).

                     Play face:Grin (teeth together, lips pulled
                     back over teeth).

  Aggressive displays:

                     Cough:Vocal threat, usually in series of three
                     or more.

                     Quadrupedal stiff stance:Animal standing, stiff
                     manner, with elbows turned inward (in tension
                     situation).

                     Stiff walk:As above, but walking quadrupedally.

                     Head divert:Head turned away from object being
                     threatened.

                     Tight lip:Lips compressed together.

                     Lunge:Animal rushing at another in short, fast
                     run (without hit).

                     Lunge with hit:As above, including a hit with
                     hand or foot.

                     Chest beat:Slapping chest alternatively with
                     hands.

                     Wall beat:As above, but directed toward
                     inanimate object in the environment (i,e.,
                     wall, pole).

                     Ground slap:Thumping the ground with one or
                     both hands.

                     Vegetation:Plucking a piece of vegetation and
                     placing it between the lips (without chewing).

                     Hand clap
                     
                     Throw vegetation:Tossing vegetation and/or rock
                     at beginning of lunge sequence.

                     Openmouth bared teeth:Lips pulled back over
                     teeth, mouth open.

  Sexual behaviors:

                     Mount:Assume sexual position.

                     Thrust:Rhythmic pelvic movements against
                     another animal.

                     Penile erection

                     Genital examination:Animal manipulating
                     another's pelvic region with fingers and/or
                     mouth.

                     Present:Animal turning its hindquarters toward
                     another.

  Other behaviors:

                     Manipulate:That is, rub, rock, pat, scratch,
                     turn, rock, bounce, etc., behavior directed
                     toward another.

                     Groom:Picking through another's hair with
                     fingers or lips.

                     Object examination:Examining objects in the
                     environment in any way except stare.

                     Social examination:Examining another with
                     fingers, feet, mouth, includes stare.

                     Confiscate food:Take food from another.

                     Confiscate object:Take object from another.

                     Confiscate infant:Animal gaining control of
                     infant from another (original animal may still
                     be in contact with infant).

                     Mouth:Excludes biting.

  Stereotypy:        Repetitive motions or vocalizations, or
                     abnormal maintenance of a posture, for which
                     there is no obvious biological significance.
.Meder, A.  (1985).  Integration of hand-reared gorilla infants in a
                    group.  Zoo Biology, 4, 1-12.

  Purpose:           To document the introduction and integration
                     processes of hand-reared gorilla infants into a
                     group.

  Sampling:          Period I: ad lib sampling
                     Period II: focal animal sampling for 5 hours
                     per 4 weeks.
                     Distances sampled at 5 minute intervals. 
                     Contacts were noted.

  Ethogram key words:Development, Introductions/Socialization.

  Behaviors:         (following Meder's dissertation, 1982)

  Stand facing:      Stand in front of partner quadrupedally and
                     look at him directly for several seconds.

  Display:           Specific behavior including stiff stance,
                     piloerection and tense lips, mostly combined
                     with further elements mentioned in Schaller
                     (1963) and Meder (1982).

  Smell:             Bring the nose very close to partner's body for
                     at least one second.

  Genital investigation:Touch ano-genital region of partner with
                     finger or lips or smell it.

  Touch with hand:   Extend one hand toward partner and touch him
                     briefly.

  Touch with body:   Sideswipe partner while moving past him.

  Run toward:        Run in the direction of partner from a
                     distance.

  Run after:         Follow partner for several meters while
                     running.

  Hit:               Slap partner with one hand once.

  Beat:              Slap partner with both hands alternately.

  Push:              Press one hand against partner briefly.
                     
  Tear:              Grasp partner and pull him briefly into
                     oneself's direction.

  Play:              Nonaggressive interaction with more than 3 s of
                     body contact and both partners participating
                     actively with such patterns as hit, push,
                     tear, hold, bump against partner, play-face
                     (mouth open widely, teeth not visible).

  Back-ride:         Rest on the back of partner, who is either 
                     standing quadrupedally or walking or climbing,
                     for several seconds.

.Meder, A.  (1986).  Physical and activity changes associated with
                    pregnancy in captive lowland gorillas (Gorilla
                    gorilla gorilla).  American Journal of
                    Primatology, 11, 111-116.

  Purpose:           To quantitatively determine changes in social
                     behavior and locomotor activity in 5 lowland
                     gorillas.

  Sampling methods:  Frankfurt:
                     Whole group observed ad lib, as well as focal
                     animal sampling. 

                     Stuttgart:
                     Focal animal sampling, nine 5 minute samples
                     per female, per observation block.
  
  Ethogram key words:Activity budgets/General behaviors, Social
behavior, Pregnancy, Behavior during.

  Behaviors:         (following Meder's dissertation, 1982)

  Long social play:  Nonaggressive interaction with at least 10
                     seconds of body contact and with both partners
                     participating actively.  Behavior patterns
                     included hitting, pushing, tearing, holding,
                     and bumping against the partner.  The play face
                     (mouth open widely, teeth generally not
                     visible) was exhibited frequently.

  Threat:            Noncontact aggression consisting of a display
                     directed toward the partner: staring at him or
                     following him while showing the display posture
                     (stiff stance, piloerection, tense lips).

  Social contact:    Behaviors directed toward a group member by the
                     focal animal.  This included all forms of
                     physical contact as well as threat.  A new
                     contact was noted when at least three seconds
                     had elapsed since the last contact to the same
                     partner had been broken.

  Locomotion:        A move of the focal animal from one place to
                     another with at least three steps.  A new
                     locomotion was noted when the female had rested
                     in one place for at least ten seconds.
.Meder, A.  (1990).  Sex differences in the behaviour of immature
                    captive lowland gorillas.  Primates, 31, 51-63.

  Purpose:           To investigate behavioural development of
                     immature gorillas and differences in terms of
                     sex and different rearing conditions. 

  Sampling methods:  Scan sampling (1 minute intervals) was used for
                     all solitary and some social behaviors.  All
                     social behaviors were observed with continuous
                     focal animal sampling.  One observation block
                     consisted of 10 hours of scan sampling
distributed over the day and 5 hr of focal animal
                     sampling in the afternoon within one week. 

  Ethogram key words:Activity budgets/General behaviors, Development.
  
  Behaviors: (not defined)(following Meder's dissertation, 1982)

  115 behavior patterns were included, including:

  Behavior of subject toward objects
  Behavior toward own body
  Behavior toward conspecifics
  Behavior toward visitors
  Locomotion
  Body movements
  Vocalizations
  Location
  Asleep
  Nest building
  Allogrooming
  Self-grooming
  Aggressive behavior:Biting, slapping, pushing, pulling.
  Social contact
  Play

.Mitchell, R. W.  (1989).  Functions and social consequences of
                         infant-adult male interaction in a captive
                         group of lowland gorillas (Gorilla gorilla
                         gorilla).  Zoo Biology, 8, 125-137.

  Purpose:           To investigate the behaviors of captive male
                     lowland gorillas to infant, of the infant to
                     the males, and of the infant's mother to the
                     behavior of the males and infant.

  Sampling:          Ad lib.

  Ethogram key words:Development, Paternal behavior.

Behaviors:

Simple actions:      The majority of simple actions described have
been "used in
                     other studies as undefined elements in
                     definitions of higher order action patterns, or
                     were presented without definition, so do not
                     require definition here" - see Estrada, 1984,
                     Tilford & Nadler, 1978.

  Combinations of simple actions:

                     Dragging:When an animal grabbed an infant in
                     contact with its mother and ran while holding
                     onto the infant, thereby pulling both the
                     infant and its mother behind it.

                     Collecting:When an animal pulled or pushed,
                     toward itself, an infant.

                     Kidnapping:When an animal moved away with an
                     infant when the kidnapper was not the primary
                     caregiver.

                     Caressing:When an animal gently moved its hand
                     around the head of another animal.

                     Cuddling:When an animal embraced another
                     whereby at least one of the animals involved
                     had his or her arms around the other.

                     Episodes:A sequence of social actions that
                     occurs when one organism performs temporally
                     close social actions toward another organism. 

.Nadler, R. D., Collins, D. C., Miller, C., & Graham, C. E.  (1983). 
Menstrual
     cycle patterns of hormones and sexual behavior in gorillas. 
     Hormones and Behavior, 17, 1-17. 

  Purpose:           To investigate possibility that different
                     aspects of sexual activity might be
                     differentially associated with two hormone
                     peaks.

  Sampling methods:  30 minute daily tests (midmorning and
                     midafternoon) of sexual behavior initiated on
                     the day after the first day of menses and for
                     each cycle continued to the next period of
                     menses.  Tests consisted of introducing female
                     with male.  

  Ethogram key words:Sexual behavior.

  Behaviors:

  Copulation:        Genital-genital contact, dorso-ventral or
                     ventro-ventral, accompanied by a series of
                     rhythmic pelvic thrusts executed by the male. 

  Male solicitations:Male approaches female, followed by male
                     pulling female toward him, preparatory to
                     covering the female.

  Covering:          Male, standing quadrupedally, assumes position
                     behind and above the female so as to
                     approximate their genitals, with or without
                     grasping the female's sides or hips, generally
                     in the dorso-ventral position.

  Male display:      Charge directed at or parallel to the female,
                     with or without prior chest- or wall-beating
                     and accompanied or not by hitting or kicking
                     the female.

  Male success:      Male solicitation that is followed by female
                     presenting.

  Female presenting: Quadrupedal stance, ranging from legs extended
                     to crouching, in which female's genitals are
                     directed toward the male.

  Male success index (MSI):Proportion of days on which male
                     solicitations were successful, multiplied by
                     100.

  Female solicitation:Female approaches the male, followed by
                     presenting, but not preceded by male
                     solicitation or male display.

  Female success:    Female solicitation followed by copulation.

  Female success index (FSI):Same but for females.

  Attractivity, proceptivity, and receptivity: [Defined as according
                     to Beach (1976)]. 

                     Attractivity: Frequency of male solicitation
                     days. 

                     Proceptivity: Frequency of female solicitation
                     days. 

                     Receptivity: MSI.
.Nadler, R. D. (1985).  Sex-related behavior of immature wild
                         mountain gorillas.  Developmental
                         Psychobiology, 19, 125-137.

  Purpose:           Providing data on sex-related behavior of
                     immature wild mountain gorillas, to supplement
                     Schaller's work.

  Sampling methods:  Ad lib sampling incidentally to a study of
                     mother-infant relations.  All observed
                     sex-related behaviors were recorded, including
                     identity of participants, positions of bodies,
                     # of pelvic thrusts, other repetitive acts of
                     genital stimulation or the duration of such
                     behavior, and group activity at time.

  Ethogram key words:Sexual behavior, Field study, Gorilla g.
beringei. 


  Behaviors:         

  Positions:         Dorso-ventral or ventral-ventral.

  Genital stimulation:Principally when animal executed thrusts
                     against the body of another, included all
                     positions.  Episode defined as a single bout or
                     multiple bouts in which bouts occurred within a
                     few minutes of each other.

  Thrusting:         Rhythmic pelvic movements.
. Ogden, J. J., Hoff, M., & Maple, T. L.  Zoo Atlanta.

  Purpose:           To study enclosure utilization and general
patterns
                     of adaptation to new enclosures in lowland
                     gorillas.

  Sampling method:   15 minute continuous focal samples of all
                     behaviors, with location scans at 15 minute
                     intervals.

  Ethogram key words:Activity budgets/General behaviors, Social
behavior, Sexual behavior, Development, Maternal behavior,
Display/Agonistic behavior, Abnormal/Stereotypic behavior.

  Behaviors:

  Solitary behaviors [no actor/recipient]:

                     Stand:Includes bipedal and quadrupedal (note
                     bipedal in
                     comments if possible).  (back legs are both
                     straight).

                     Locomote:Any moving of the animal's entire body
                     including walking, running [bipedal or quad].

                     Sit:Includes squat and leaning on one elbow.

                     Lie: Dorsally, ventrally, or on side (includes
                     lying with one shoulder touching ground, or
                     leaning ventrally on both  elbows).

                     Climb

                     Autogroom:Pick through own hair with fingers
                     and/or lips.

                     Masturbation
                     
                     Solitary play:Self-motion play.

                     Feed/forage:Manipulating the substrate or food
                     item while intermittently transferring items to
                     the mouth or chewing a food item.  Note item
                     eating (i.e., browse, monkey chow, fruit, tree
                     branch).  Includes drinking, but note if
                     drinking from stream or lixit.

                     Object manipulate:The handling of available
                     objects with hands, feet, or mouthing;
                     examining except stare.  [Note: score this
                     category if sorting through straw or
                     manipulating branches/browse without movement
                     to mouth].  Note object being manipulated.

                     Carry/hold:The carrying or holding of object
                     with hands or feet.   Either while locomoting
                     or while stationary.  (Or carrying on back
                     while walking). Note object.

 .Social Behaviors - note actor/recipient:

                     Approach:Approach another to within one arm's
                     length.

                     Displace:Take place of other.

                     Move away:Move away from another, either from
                     within contact or within arm's length.

                     In contact:In contact with another, any
                     posture.  Note posture.

                     Sexual behavior:Includes mount, insertion,
                     genital inspection, etc.  Note in comment
                     section.

                     Social playMotion play; social play involving
                     object manipulation should be noted as SP/OM.

                     Groom-social:Pick through hair of other with
                     fingers and/or lips (excludes scratching).

  Displays/Threats/Vocalizations: (note actor/recip where  known]

                     Rumble/belch vocal.

                     Cough vocalization:Vocal threat; usually in
                     series of three or more.

                     Quad. stiff stance:Animal standing; stiff
                     manner; with elbows turned inward (in tension
                     situation).

                     Stiff walk:As above, but walking quadrupedally.

                     Tight lip:Lips compressed together.

                     Openmouth bared teeth:Lips pulled back over
                     teeth, mouth open.

                     Agonism:Includes biting, fighting, hitting,
                     slapping, lunge with hit; any aggression with
                     contact.  Time interval between bouts of 15
                     seconds.

                     Lunge:Animal rushing at another in short, fast
                     run (without hit). 

                     Chest beat:Slapping chest alternately with
                     hands (includes beating of thighs by females).

                     Wall beat:Beat/slap directed toward inanimate
                     object in environment (note object).

                     Ground slap:Thumping ground with one or both
                     hands.

                     Hand clap

                     Throw objectTossing object at beginning of
                     lunge sequence.

. Abnormal Behavior (non-social; no actors/recip):

                     Self-biting:Usually of arms, legs, torso,
                     hands.

                     Self-clasping/Embracing, or use of hands or
                     feet to
                     self-grasping:hold onto a part of body.

                     Stereotyped pacing

                     Rocking:Often associated with self-clasping.

                     Coprophagia:Ingestion of fecal material (not
                     necessarily their own).

                     Regurgitation/
                     reingestion:Regurgitation and reingestion of
                     feeding matter.

                     Out of viewNote substrate if known, tunnel if
                     known.

                     OtherAny behavior of interest not listed here;
                     explain in comments.

  Maternal/infant behavior:

                     Active social play:Whole body movement,
                     including rough wrestling, rolling, chasing.

                     Passive social play:Less than whole body
                     movement, often gentle rocking, mouthing or
                     wrestling.

                     Initiate play:Animal responsible for beginning
                     social play bout.

                     Retrieve:Retrieve another while moving away.

                     Remove:Move another into a lesser contact
                     (e.g., VV to OC).

                     Restrain:Prevent another from moving away.

                     Wean:Remove infant from nipple.

                     Social examination:Examining another with
                     fingers, feet, mouth; includes stare.

                     Nipple contact

                     Manipulate:Rub, rock, pat, scratch, turn, rock,
                     bounce, etc.; behavior directed toward other.

                     Self-manipulation:As manipulate only
                     self-directed.

                     Struggle:Infant attempts to leave while Mother
                     is restraining.

. Contact Variables:

                     Ventro-ventral:Common usage.

                     Dorso-ventral:M dorsal, I ventral; i.e., back
                     ride position.

                     Ventral-dorso:As above, but vice-versa.

                     Other contact:Any contact other than above or
                     "touch".

                     Proximate:Out of contact, within arm's length.

                     Distant:Greater than arm's length but less than
                     15 feet.

                     Far distant:Greater than 15 feet from another
                     animal.

  Maternal Transport:

                     Tripedal walk:Mother holding infant with one
                     arm while walking.

                     Hanging walk:Animals VV, mother walking
                     quadrupedally.

                     Extended arm drag/carry: Mother carrying infant
                     away from her body and along/or off the ground.

                     Back ride:Infant riding on mother's back,
                     "jockey style".

                     Crutch walk:Infant in mother's lap, M move
                     using arms as crutches.

                     Other transport
                     positions

.Quiatt, D., Miller, L., & Cambre, R.  (1986).  Overt behavior
correlates
                                             of menstruation and
                                             ovulation in a lowland
                                             gorilla.  In D. M. Taub
                                             and F. King (Ed.),
                                             Current perspectives in
                                             primate biology (pp.
                                             32-41).  New York: Van
                                             Nostrand Reinhold.

  Purpose:           To investigate whether observations of social
                     behavior might help pinpoint ovulation
                     sufficiently to increase chances of successful
                     artificial insemination.

  Sampling:          Eight weeks of data collection, during each of
                     two summers.  Most days 1 hour of observation
                     in the morning and 1 in the afternoon,
                     scheduled when gorillas were most active
                     (during 2 hours following morning feeding and 2
                     hours preceding afternoon feeding). 
                     Frequencies/rates resulted. 

  Ethogram key words:Social behavior, Sexual behavior.

  Behaviors:

  Displacement:      One individual "causes" another to move away
                     from the location he/she has been occupying. 
                     The first may or may not replace the second in
                     space.  Cause may be assumed from apparent
                     intent of the displacer (e.g., gaze or eye
                     aversion) or attentiveness of the displaced
                     individual.  Rather than infer cause, and to
                     insure consistency, displacement recorded as
                     whole body movement of one individual which was
                     one or more feet away from the other, when the
                     other is in movement and has approached within
                     six feet.

  Proximity:         One individual moves to within arm's reach of
                     another and remains without other interaction
                     for at least 5 seconds.  If the approached
                     individual moved off before 5 seconds was up,
                     we scored the behavior as "Displacement." 
                     (Note the qualifier, "without other
                     interaction."  This is what distinguished
                     "Proximity" as an observational unit.)

  Contact:           Any tactile communication other than
                     "Examination," e.g., touching, brushing,
                     hitting, sitting, lying in contact, etc.

  Groom:             No grooming of either animal occurred, so
                     eliminated this.

  Examination:       An inspection of one animal by another or a
                     self-inspection.  Examinations involved
                     looking, touching, smelling, mouthing, and
                     combinations thereof. 

  Display:           A stereotypical hammering of the chest with the
                     knuckles.  We noted two levels of intensity: 
                     (1) a low level, without accompanying
                     locomotion or other signs of agitation - 1 or 2
                     taps or a series of light taps (the displaying
                     individual might be sitting or lying), and (2)
                     a rapid series of forceful chest-pounding while
                     the displaying individual was standing and
                     perhaps locomoting.  Many such displays were
                     directed toward spectators.  If a display
                     involved contact with the window, we noted that
                     contact.
.Riess, B. F., Ross, S., Lyerly, S. B., & Birch, H. G.  (1949).  The
     behavior of two captive specimens of the lowland gorilla
     Gorilla gorilla gorilla (Savage & Wyman).  Zoologica, 111-118.

  Purpose:           To provide basic information on lowland
                     gorillas, comparable to that of Carpenter's.

  Sampling:          During three-minute periods of observation all
                     behaviors which occurred were checked.  30
                     minute observation periods.  Sequences were
                     noted.

  Ethogram key words:Activity budgets/General behaviors, Social
behavior.

  Behaviors:

  42 behaviors divided into the following five categories

  Posture and locomotion:

                     Walking, pronograde
                     Walking, upright
                     Running, pronograde
                     Running, upright
                     Standing, pronograde
                     Standing, upright
                     Sitting
                     Climbing
                     Swinging by hands
                     Hanging
                     Sliding
                     Left-handedness
                     Right-handedness
                     Lying down
                     Sleeping
                     Nest building

  Eating, drinking and elimination:

                     Gathering food in heaps
                     Sharing food with cage mate
                     Drinking water
                     Drinking milk
                     Regurgitation of milk
                     Handling of feces
                     Attention to urination

  Self-oriented activity:

                     Self-manipulation (non-genital)
                     Manipulation of genitalia
                     Manipulation of objects
                     Manipulation of food (non-eating)
                     Attitude toward stream of water
                     Self-grooming

. Inter-individual behavior:

                     Chasing
                     Wrestling
                     Grooming
                     Inspection and manipulation of genitalia
                     Presenting
                     Mounting
                     Pelvic thrusts
                     Chest thumping
                     Vocalizing
                     Dominance

  Observer-oriented behavior:

                     Throwing of feces
                     Throwing of non-fecal material
                     Attentional responses
                     Vocalization
.Schaller, G.  (1963).  The mountain gorilla: Ecology and behavior. 
                         Chicago: University of Chicago Press.

  Purpose:           To provide the first systematic examination of
                     behavior of wild mountain gorillas. 

  Sampling methods:  Ad lib field notes.  

  Ethogram key words:Activity budgets/General behaviors, Sexual
behavior, Social behavior, Display/Agonistic behavior,Field study,
Gorilla g. beringei.

  Behaviors:A separate ethogram is not provided, but behavioral
descriptions
                     are integrated into the text.  The reader is
                     referred to the book.
. Schildkraut, D. & Akers, J.  Boston MetroParks Zoos.

  Purpose:           To compile baseline and comparison data on a
                     pair of gorillas before and after their move
                     from Cincinnati Zoo to Stone Zoo.

  Sampling methods:  All occurrences.

  Ethogram key words:Activity budgets/General behaviors, Social
behavior.

  Behaviors:

  Social behaviors:

                     Play:Wrestling, chasing, running past and
                     cuffing each other, gentle slapping and
                     grabbing, bipedal jostling (time + frequency). 
                     Initiator listed first.  Note: a  play bout
                     must be separated by a 10 second interval or
                     another behavior to constitute another
                     frequency.

                     Groom:Animal mouths and manipulates the fur of
                     another.

                     Touch:Animals in contact when not engaged in
                     any other defined activity (time + frequency). 
                     Initiator listed first.

                     Arm's length:Animal is within arm's reach of
                     another and not engaged in any other defined
                     activity  (time + frequency).  Initiator listed
                     first.

                     Displace:One animal approaches another, and the
                     2nd animal moves away before they are within
                     arm's reach (frequency).  Initiator listed
                     first.

                     Fight:Rough hitting or biting accompanied by
                     screaming.

  Self Behaviors:

                     Display:A single incidence of banging/kicking
                     doors and walls OR a series of events which
                     includes chest beating, stamping feet, running
                     and pulling chain, loud clapping.  The display
                     is usually performed bipedally and ends in
                     running a short distance (frequency).

                     Self-groom:One animal mouths and manipulates
                     its own fur for more than 5 seconds
                     (frequency).  Note:  self-groom does NOT
                     include licking fingers during the process of
                     regurgitation and reingestion.

.                    Self-play:Sliding on the floor when wet,
                     turning in circles on all fours, clapping
                     softly and shaking head or limbs, rolling on
                     back and slapping feet with hands, hanging
                     upside down from chain or ladder, squirting
                     self with water spout, grabbing hose from
                     keeper, putting food on head and "laughing",
                     swinging on ladder, somersaults, spitting into
                     hands or on floor,  pushing food along the
                     floor, wiggling chain, standing on hands and
                     walking up the wall, sticking out tongue,
                     pushing food out of cage, self-tickling.  AW =
                     a behavior typical of the female where she
                     scoots along the floor on her rear end for
                     short distances.  The female also puts her
                     knuckles or fingers in her eyes and walks about
                     the cage "as if she were blind" (frequency).

                     Regurgitation andFood or fecal material is
                     eaten, regurgitated back into the
                     reingestion:hands or floor and eaten again
                     (frequency).

                     Cage change:Animal moves from cage I to cage II
                     through doorway  (frequency).
. Schildkraut, D. (a), Boston MetroParks Zoos.

  Purpose:           To determine the dominance relationship between
                     an adult male and female pair of gorillas, with
                     particular regard to its possible effect on
                     their breeding behavior.

  Sampling methods:  Frequency and sequential recording, three
                     ten-minute intervals per day.

  Ethogram key words:Activity budgets/General behaviors, Social
behavior, Sexual behavior.

  Behaviors:

  Gigi crosses into Sam's exhibit

  Sam sits and calls at Gigi's door

  Grooming

  Feeding/stealing/
  bribing with food

  Spatial displacement

  Aggressive behavior:Such as hitting, biting, etc.

  Threats:           Specifically directed at the other gorilla -
                     not at staff or public.

  Social encounters

  Breeding
. Schildkraut, D. (b), Boston MetroParks Zoos.


  Purpose:           Ethogram for introduction of three peer-raised
                     juvenile gorillas to a mother-infant pair.

  Sampling methods:  Frequency recording, one hour per day, five
                     days per week, over a twelve month period.

  Ethogram key words:Introductions/Socialization, Social behavior,
Development, Maternal behavior.

  Behaviors:

  Self-directed behaviors:

                     Self-grooming

                     Sleeping

                     Sliding

                     Walking:Specify bipedal or quadrupedal.

                     Swinging

                     Spinning

                     Handstands

                     Hand clapping

                     Hand shaking

                     Eating

                     Urinating

                     Defecating

                     Object manipulation:Specify the object and
                     indicate how object is manipulated, i.e.
                     thrown, banged, put on head, etc.

                     Stare

                     R & R:Regurgitation and reingestion of food.

                     Coprophagy:Eating feces.

  Noncontact social behaviors:

                     Quadrupedal straight leg stance

                     Open mouth threat

                     Chest beating

                     Bluff charges
 
                     Vocalizations

                     Approach:Move toward another animal.

                     Affiliate:Sits within one arm's length of
                     another.

                     Displace:One animal moves out of the way as
                     another approaches.

  Contact social behaviors:

                     Wrestling

                     Tumbling

                     Charging with hitting

                     Chasing

                     Tickling

                     Hitting

                     Pushing

                     Biting

                     Mounting

                     Copulating

                     Grooming others

                     Sit/touch:Sits or reclines in contact with
                     another animal.

  Mother/infant behaviors:

                     Nursing:Specify which breast (right or left).

                     Carrying:Specify v-v, v-d, d-v, arm, leg,
                     thigh, etc.

                     Retrieve:Mother retrieves infant.

                     Restrain:Mother prevents infant from leaving
                     her.

                     Struggle:Infant struggles to get away from
                     mother.

  Other:

                     Follow:May be used when any animal follows
                     after another.  Often seen in mother/infant
                     pairs.

                     Cage change:Used when introductions take place
                     to indicate that an animal has moved from one
                     cage to another.
.Schildkraut, D. (c), Boston MetroParks Zoos.

  Purpose:           Observations on the behavior of an isolated
                     adult male gorilla.

  Sampling methods:  Frequency recording, three ten-minute periods
                     per day.

  Ethogram key words:Activity budgets/General behaviors.

  Behaviors:

  Staring:           Watching or looking at something.

  Groom/body part:   Specify part of body being groomed.

  Slide on floor:    Usually occurs when cage floor is wet.

  Object manipulation
  in quiet play:     Playing with objects while in a stationary
                     position.

  Object slapping:   Beating or slapping on an object with hands or
                     feet - specify which he uses, and the object.

  Bluff charges:     Incomplete charges, specify target (i.e., zoo
                     keeper).

  Charges with hitting:Usually hits the front glass or the side
                     walls.  Specify target.

  Sitting

  Climbing

  Running

  Sleeping:          Specify on floor or platform.

  Eating

  R & R:             Regurgitation and reingestion.

  Coprophagy:        Eating feces.

  Chest beating

  Facial threat:     Usually open mouth.

  Threat vocalization

  Threat stance
. Schildkraut, D. (d), Boston MetroParks Zoos.
    
  Purpose:           Observations on the introduction of an adult
                     male/female pair.

  Sampling methods:  Positions mapped at beginning, after 5 min.,
                     and at end of observation period.  All
                     occurrences of the following behaviors were
                     recorded, noting actor and recipient.

  Ethogram key words:Introductions/Socialization.

  Behaviors:

  Full body display

  Head display:      Usually done from a sitting position.
                     
  Charge

  Charge with slam

  Lunge

  Rattle/bang door:  From a stationary position.

  Approach door:     Moving toward the door in a non-threatening
                     manner, i.e. not charging.

  Sitting at door

  Passing objects
  through door

  Physical contact:  Touching the other animal.

  Withdrawal from
  door area

  Display with chest
  beating
. Schildkraut, D. (e), Boston MetroParks Zoos.

  Purpose:           Gorilla Enclosure Utilization Checksheet.

  Sampling methods:  15 minute continuous focal samples of all
                     behaviors, with scans at 15 minute intervals of
                     locations.

  Ethogram key words:Activity budgets/General behaviors, Social
behavior, Sexual behavior, Development, Maternal behavior,
Display/Agonistic behavior, Abnormal/Stereotypic behavior.

  Behaviors:

  Same as those listed by Ogden, J. J., Hoff,  M. and Maple,  T. L.
                     with two additions:

  Diarrhea

  Ventral/ventral embrace
.Smalley, S. L.  (1979).  Comparative analysis of aggressive and other
                         social behavior of captive lowland gorillas
                         and free living mountain gorillas. 
                         California Anthropologist, 9, 1-28.

  Purpose:           To compare the interactions and behaviors of
                     captive gorillas to those of free living
                     mountain gorillas. 

  Sampling methods:  Interactions were recorded during or
                     immediately after behavioral sequences.  Group
                     was observed as a whole, interactions given
                     priority over solitary behaviors. 

  Ethogram key words:Activity budgets/General behaviors, Social
behavior, Display/Agonistic behavior, Field study, Gorilla g.
beringei.

  Behaviors:

  Aggression:        For purposes of study defined as dominance: one
                     gorilla is considered dominant over another if
                     the first displaces the second, with or without
                     physical contact. 

  Other behaviors recorded (definitions unclear):

                     Feeding
                     Chestbeating
                     Self-grooming
                     Mutual grooming
                     Object carrying/holding
                     Running
                     Rising as part of a display
                     Rubbing of underarms
                     Copulation
                     Sexual presentation
.Watts, D. P.  (1988).  Environmental influences on mountain gorilla
                         time budgets.  American Journal of
                         Primatology, 15, 195-211.

  Purpose:           To investigate mountain gorilla time budgets
                     and the effect which characteristics of their
                     food supply and other ecological factors have
                     on them, including sex differences. 

  Sampling:          Focal sampling to collect time budget data,
                     durations recorded.

  Ethogram key words:Activity budgets/General behaviors, Field study,
Gorilla g. beringei.

  Behaviors:         (See also Watts, D. P.  (1983).  Foraging
                     strategy and socioecology of mountain gorillas. 
                     Ph.D. Diss., University of Chicago.)

  Feeding:           Preparation and ingestion of food.  Feeding
                     bouts began when a stationary individual,
                     either on arrival at a feeding spot or after
                     having engaged in some other activity there for
                     at least 5 seconds, started to prepare food for
                     ingestion (which included chewing).  They ended
                     when the individual ceased preparation and
                     ingestion for at least five seconds or shifted
                     position by at least 1 m.  

  Moving:            All locomotor activity other than shifts of
                     position of less than 1 m during feeding and
                     resting and locomotion during play. 

  Resting:           Stationary, nonsocial behavior during which
                     animals were not feeding (sleeping,
                     autogrooming, etc.) 

  Social:            Social interactions with a measurable duration,
                     including play, allogrooming, sexual behavior,
                     and some forms of agonistic behavior. 

  Other:             Other subsistence behavior, including nest
                     building, and, for females, nursing.  Because
                     it was usually impossible to be certain whether
                     a female was nursing when she had her back
                     turned, and because "other" activities occupied
                     only a tiny fraction of observation time, they
                     are not discussed below.
.Watts, D. P.  (1990).  Mountain gorilla life histories, reproductive
                         competition and sociosexual behavior and
                         some implications for captive husbandry. 
                         Zoo Biology, 9, 185-200.

  Purpose:           To draw implications from the data on wild
                     mountain gorillas to apply to captive
                     management and husbandry of captive lowland
                     gorillas.

  Sampling:          All occurrences sampling of below behaviors.

  Ethogram key words:Sexual behavior, Field study, Gorilla g.
beringei.

  Behaviors:         (See also Watts, D. P.  (1983).  Foraging
                     strategy and socioecology of mountain gorillas. 
                     Ph.D. Diss., University of Chicago.)

  Sociosexual behavior:Heterosexual mounts with thrusting, with or
                     without intromission.

  Copulations:       Mounts in which one or both partners thrust and
                     positioning by both suggests that intromission
                     occurred.
. Woods, S.  Denver Zoo, Enrichment Study.

  Purpose:           Investigate effects of enrichment protocols and
                     general activities.

  Sampling:          Focal animal, behavioral checksheet.

  Ethogram key words:Enrichment, Activity budgets/General behaviors.

  Behaviors:

  Aggressive interaction - with another gorilla:

                     Display threat
                     Physical attack
                     Chase
                     Displacement
                     
  Other -- urinate, ambiguous, etc.:

                     Drink
                     Sexual activity - mating attempts, solicit
                     Playful social interaction

  No observation:

                     Not visible
                     Bad observation (cannot see behavior)

  Stereotypic or undesirable behaviors:

                     Regurgitation/reingestion
                     Coprophagy
                     Masturbation

  Self-grooming (must be manual and visual)

  Manipulate:

                     Manipulate, investigate, With objects or
                     exhibit features (takes precedence over
                       play:public orientation).
                     Manipulation of
                     enrichment items

  Travel:

                     Quadrupedal walking
                     Quadrupedal running
                     Climbing
                     Eat while traveling
                      (has priority)

. Public orientation:

                     Intently watching
                       humans
                     Display: e.g., chest beating, charging, hitting
                     glass, baring teeth.
                     Gesturing: e.g., clap, treat sign.
                     Vocalizing: (can see chest vibrate).

  Idle:

                     Sit or squat
                     Stand
                     Recline
                     Intently watching
                       another animal:(has priority)

  Feed:

                     Foraging: Searching in or under hay, eating
                     hay, seeds, nuts.
                     Foraging while intently
                       watching public
                     Grass
                     Eating from enrichment
                       objects
                     Eating anything else
                     Eating while intently
                       watching public
                     Browse

  Object contact:

                     Structural features: Walls, doorways, back
                     fence, front glass, platforms.
                     Furnishings: Trees, shelves, net, ropes, log,
                     rocks, poles.
                     Enrichment objects
                     Other portable objects

  Outside:           Includes doorway to outside yard.

  Up:                On shelves, trees, net.
.Wood, G., Forthman, D. L., & Ogden, J. J.  Ethogram for enrichment
                                             study, Zoo Atlanta.

  Purpose:           To study general activity budgets with a focus
on
                     those behaviors likely to be affected by
                     presentation of enrichment items. 

  Sampling method:   Group scans at 2 minute intervals, for hour long
                     observation periods.

  Ethogram key words:Enrichment, Activity budgets/General behaviors,
Social behavior, Display/Agonistic behavior.

  Behaviors:

  Position and activity:

                     Lie: Dorsally, ventrally, or side (includes
                     lying with 1 shoulder touching ground, or
                     leaning ventrally on both elbows).

                     Sit:Includes squat and leaning on one elbow.

                     Stand:Includes bipedal and quadrupedal  (back
                     legs are both straight).

                     Locomote:Any moving of the animal's entire body
                     including walking, running [bipedal or quad].


                     Climb:Climbing a substrate.

                     Hang/swing:Hanging or swinging from a
                     substrate.

  Solitary behaviors:

                     Autogroom:Pick through own hair with fingers
                     and/or lips.

                     Object manipulate:Manipulating available
                     non-food item with hands, feet, or mouth: e.g.,
                     branches, rocks, or bark (if not attempting to
                     eat such items).  

                     Carry/hold:Carrying or holding a non food item
                     with hands, feet, or draped across back.  
                     Either while locomoting or stationary. 

                     Solitary play:Self-motion play.

                     Masturbation:Manipulating genitals.
                     
  Social Behaviors - note actor/recipient:

                     Proximity:Within one arm's length to another
                     group member.

                     Social contact:In contact with another group
                     member.

                     Social groom:Pick through another's hair with
                     fingers or lips.

                     Social play:Includes all motion social play
                     behaviors: chase, somersault, hit, etc.

                     Sex:Includes mount, insertion, genital
                     inspection, etc.

  Displays/Threats:

                     Quad. stiff stance:Animal standing; stiff
                     manner; with elbows turned inward (in tense
                     situation).

                     Stiff walk:As above, but walking quadrupedally.

                     Tight lip:Lips compressed together.

                     Bared teeth/openmouth:Mouth open with lips
                     pulled back over teeth.
                     Agonism:Aggression with contact, including:
                     biting, fighting, hitting, slapping, and lunge
                     with hit.

                     Lunge:Animal rushes at another (without hit). 

                     Chest beat:Slapping chest alternately with
                     hands (includes beating of thighs by females).

                     Wall beat:Beat/slap wall or other substrate.

                     Throw object:Throwing an object, frequently
                     exhibited before lunge.

  Abnormal Behavior:

                     Self-biting:Usually of arms, legs, torso,
                     hands.

                     Self-clasping/Self-embracing, or holding onto a
                     body part.
                     self-grasping:

                     Stereotyped pacing:Pacing, walking in circles.

                     Rocking:Often associated with self-clasping.

                     Coprophagia:Ingestion of any fecal material.

                     Regurgitation/
                     reingestion:Regurgitation and reingestion of
                     feeding matter.

                     Out of view:Note substrate if known, tunnel if
                     known.

                     Other:Any other behavior of interest not
                     listed.

  Feeding behavior:

                     Carry/hold food:Carry or holding food or
                     enrichment item.  Note item.

                     Foraging:Gathering and collecting food items. 
                     Note item.

                     Process food:Processing, preparing, stripping,
                     peeling, seek, remove pith, sucking, husking,
                     and chewing.  Some processing may take place
                     inside the buccal cavity (e.g., peeling).  Note
                     item.
Yamagiwa, J.  (1986).  Activity rhythm and the ranging of a solitary
                         male mountain gorilla (Gorilla gorilla
                         beringei).  Primates, 27, 273-282.

  Purpose:           Description of activities, ranging behavior and
                     responses to neighboring groups by a lone male
                     gorilla.

  Sampling
  (of behavioral info):  Focal sampling, each activity was scored in
                     every 10 minutes by 10 second observations.

  Ethogram key words:Activity budgets/General behaviors, Field study,
Gorilla g. beringei.

  Behaviors:

  Feeding

  No move

  Locomotion

  Others:

                     Chest-beating display?
                                 .INDEX

Abnormal/Stereotypic behavior 7, 27, 48, 54, 92, 107

Activity budgets/General behaviors 3, 7, 25, 27, 37, 50, 54, 73, 79,
81, 87, 88, 92, 97, 99, 100, 102, 105, 107, 108, 109, 113, 115

Birthwatch 46

Development 5, 27, 41, 52, 70, 72, 73, 81, 86, 88, 89, 92, 103, 107

Display/Agonistic behavior 7, 27, 38, 54, 73, 81, 92, 99, 107, 108,
113

Enrichment 111, 113

Environment change, Influence of 25, 48

Field study1 37, 39, 41, 45, 67, 68, 69, 91, 99, 108, 109, 110, 115

Gorilla g. beringei 37, 39, 41, 45, 67, 68, 69, 91, 99, 108, 109,
110, 115

Introductions/Socialization 38, 54, 86, 103, 106

Maternal behavior 27, 73, 81, 92, 103, 107

Noise disturbance, Effects of 50

Paternal behavior 89

Post-occupancy evaluation 7, 48

Pregnancy, Behavior during 41, 87

Sexual behavior 7, 27, 45, 54, 73, 81, 90, 91, 92, 96, 99, 102, 107,
110

Social behavior 3, 7, 26, 27, 54, 67, 68, 69, 73, 81, 87, 92, 96,
97, 99, 100, 102, 103, 107, 108, 113

Vocalizations 27, 39, 54, 73






1 Studies not designated as field studies were conducted in
captivity.