Giant Panda Online - Species Survival Plan

Giant Panda Behavior Research Methods
Dr. Ron Swaisgood, C.R.E.S

In this section an overview of the behavior research methods used by San Diego Zoo research staff is presented. The version here is the one used in Wolong, which differs only slightly from the one used in San Diego. We have found that each institution may have slight variations (for example, enclosure design, access to conspecifics) that requires minor modifications to the methodology to accommodate each unique situation. The version offered here is also a somewhat abbreviated version of our working methodology. In the longer version, we provide details of the minutiae and nuances regarding what constitutes a particular behavior, and supplemental rules that guide the observer through many decisions that need to be made in specific—and often rare—situations. Still, the version presented here is quite long and comprehensive, for it meets the needs of more than a dozen different topics subject to study by our research team. We recommend that anyone wishing to use this methodology should pick and choose behaviors and protocols tailored to their own specific research needs.

The observer is provided with a detailed ethogram (a list of behaviors with precise operational definitions), a recording protocol, a behavior data collection sheet, and a data summary sheet.

Behavior data collection sheet.

A place to record standard “header” information is found at the top of the data collection sheet. Here we record the subject’s identity, the date and time, the observer’s name, the enclosure, the temperature and cloud cover, the identity of neighboring pandas, and the number of visitors present. In addition to this standard information, we record an estimate of the quantity and quality of bamboo available during the session. Bamboo availability is rated by the following criteria on a scale from 0 to 3 at intervals of 0.5.

Bamboo quality index:

0. No bamboo present or available bamboo is in very poor condition and most of preferred portions are already eaten. Often recorded for leftovers present in the morning prior to the day’s first feeding.

1. Panda is given a small amount of fresh bamboo (e.g., less than 25% of a “typical” feed or there is little bamboo remaining from a previous feeding and most is old and of poor quality, yet some “desirable” portions remain).

2. Ample relatively fresh bamboo remaining, although some may have been eaten earlier.

3. Ample fresh bamboo present, placed in the enclosure just prior to or during the observation period.

Note: if the status of bamboo changes during the observation session, calculate an average bamboo quality index for the session based on the length of time bamboo of a particular rating was present. For example, if in a 45-min session, a “0” is scored for the first 30 min and a “3” is scored for the last 15 min, the average score is: (0+0+3)/3= 1.

Below the header are 30 rows for collection of behavioral data. Each row represents one minute. The first column is for recording all occurrences of behaviors occurring during the one-minute interval. The second column is for recording of vocalizations using the “one-zero” method. The next column is for recording the “point sample” behavior at the end of each one-minute interval. In the last three columns the panda’s location, social distance to other pandas, and “interaction potential” with other pandas are recorded. For definitions see below.

Behavior Data Recording Protocol

Terminology for behavioral categories.

• STATES (S) are generally behaviors that are long in duration, such as resting, feeding, or locomoting. We only record STATES in all-occurences if the behavior lasts more than 5 seconds (see below).

• EVENTS (E) are generally behaviors that are of relatively short duration, such as somersault, head-toss, and approach.

• STATE-LIKE EVENTS (SLE). Not all behaviors fall neatly into these two categories. Some EVENTS can occur for more than five seconds, making them "STATE-LIKE". For example, a somersault only takes approx. 1-2 seconds, so it would qualify as an EVENT. However, if a panda somersaults down a hill, it could take 7-8 seconds. Therefore, the EVENT "somersault" now becomes a STATE of somersaulting which is longer in duration. These "STATE-LIKE EVENTS" do not have to occur for five seconds before they are recorded because we treat them as EVENTS; however, in all other respects, they are treated like STATES.

We use three fundamental behavior recording methods:

All-occurrence sampling. This method is used to record each instance of a behavior as it occurs, and provides an accurate record of the frequency of the behavior. A behavior may be recorded several times within a one-minute interval. Each EVENT is recorded every time it is observed, regardless of how long the event lasts or how much time passes between each occurrence of the same event. STATES must last for > 5 seconds before it is recorded. A particular state must be interrupted for more than 5 seconds before it is recorded a second time. For example, if the panda locomotes for 8 seconds, rests for 3 seconds, then continues to locomote, only a single bout of locomotion is recorded and rest is not recorded at all. SLE’s need not last 5 seconds, but they must be interrupted for > 5 seconds before another bout of the same behavior is recorded.

One-zero sampling. This method is most useful for recording behaviors that occur too frequently to record each instance (e.g., vocalizations, tongue-flicking) or for behaviors in which the onset and termination are difficult to identify (e.g., tail-up and many forms of play). If a behavior designated as a one-zero behavior occurs, the behavior is recorded once and only once during that one-minute interval. If it is observed several times during an interval, it is still recorded only once. If it is not observed during that interval, it is not recorded. Thus, it is recorded one or zero times per interval, hence the label “one-zero” sampling. Note that if a behavior occurs at the end of one interval and carries over into the next interval, it is recorded in both intervals: the behavior need not stop and start as in all-occurrence sampling. This method of sampling yields a type of frequency, but not the “true” frequency provided in all-occurrence sampling. Only behaviors designated as “1-0” in the ethogram are recorded in this manner.

Point-in-time sampling. This method is used to estimate the percent time that animals spend in various STATES or STATE-LIKE EVENTS. At the end of each one-minute interval a “beep” will sound, and the observer records the STATE or SLE that the animal is engaged in at precisely the time of the beep. Percent time is calculated by dividing the number of intervals the behavior was observed at the “point sample” by the total number of intervals during the observation session (e.g., 30/100=30%). A hierarchy for which behavior to record when more than one behavior is observed at the beep is provided. In general, only STATES and SLE’s are recorded at the point sample. However, a few EVENTS and 1-0 behaviors are also recorded at the beep, designated by “at beep” in the ethogram. These are behaviors that we wish to record frequencies using one-zero or event protocols, but wish to obtain a percent time estimate for these behaviors at the point sample as well. A good example is the stereotypic behavior “head toss.” Most pandas generally head-toss in a very event-like way, e.g,, once will turning a corner. A few pandas, however, spend a great deal of time head tossing, and may stop and head toss 10-15 times in a row. In this case, we’d like to know how many head tosses were observed (i.e., as events), but we’d also like to know how much time they spend head-tossing (point sample). Note that since true events are very short in duration (e.g., a vocalization, though recorded using 1-0 method) an animal is usually engaged in some behavioral state (e.g., locomotion) simultaneously. Thus, if an animal is locomoting and vocalizing at the beep, locomotion is recorded. Note that the behavior data sheet also has columns at the end of each interval to record other information at the point sample, such as location and social distance.

Giant Panda Ethogram, Abbreviated Version

This ethogram is the product of several years of intensive observations of giant pandas. The original version was based largely on published accounts of panda behavior available at the time from the work of Devra Kleiman and George Schaller. Initially, we used a recording system in which each behavior was identified by a numeric code. Often we found that our research needs required that these general categories be divided into several subcategories, and we instituted the practice of identifying subcategories by following the numeric code with a letter (e.g., feed on bamboo = 1a). As time passed we began to see many behaviors that had not been reported before, and we added these to the ethogram. Panda behavior can be very idiosyncratic (especially stereotypies) and as we increased the number of pandas observed in China, we added new behaviors to describe what we observed. We also greatly expanded the scope of our research, and with each new project we added numerous new behaviors to meet the needs of the research question. For example, we now have ongoing research programs on chemical communication, acoustic communication, several aspects of reproductive behavior, social behavior, pregnancy monitoring, mother-infant relationship, behavioral development, psychological well-being, environmental enrichment, cognition, color discrimination, and bamboo feeding. Eventually the numeric system grew unwieldy, and as we added new behaviors, we used a shorthand code that used 2 or more letters from the behavioral descriptor (e.g., BO = Bite Object, AP = approach). The resulting ethogram, therefore, reflects this evolutionary history.

GIANT PANDA ETHOGRAM, SAN DIEGO

Miscellaneous Individual Behaviors.

Code	S-E	Activity	Definition
0	S	Not visible	Animal moves temporarily out of view. (Score for point samples only.)
1a	S	Feed on bamboo	Feeding on provisionedbamboo. Includes both consumption (chewing) and handling of food, for example, peeling bamboo stalks, stripping leaves, or any other manipulation of bamboo to prepare it for consumption.
1b	S	Feed on provisioned food	Feeding on any other provisioned food (bread, vegetables, fruit). Feeding defined as in 1a above.
1c	S	Forage on bamboo	Feeding on bamboo growing in enclosures. Feeding defined as in 1a above.
1d	S	Forage- other	Feeding on any vegetation growing in enclosures. Note food type. Feeding defined as in 1a above.
1E	S	Feed on enrichment	Feeding on an enrichment item not specifically provided for that purpose (e.g. ice, hay, bamboo puzzle feeders). Feeding defined as in 1a above.
FS	S	Food Search	Any behaviors associated with looking for biscuits, such as walking slowly while sniffing the ground, or sitting and sniffing the area for more biscuits. FS may only be scored within the context of a feeding bout and includes re-visiting sites where food has been offered in the past. FS also includes climbing to look for food or standing bipedally to sniff the structure. FS need not be followed by feeding. EFS is scored when the panda searches for food items that have been removed from enrichment itemssuch as puzzle feeders. Use of the paws should be scored as PAW (see below).
2	S	Locomote	Short bout of directional travel between points, or sustained locomotion in a non-stereotyped manner. If panda walks through waterplease note substrate and location (i.e. 'g-drinker' or 'g-pond') each time panda walks through water during the locomotion bout (i.e. '2-g-drinker, -g-pond, -g-pond). Walking through water does notinterrupt a locomotion bout (i.e. treat as a true event).
3	S	Stereotypic Pacing	Back and forth, or perimeter travel in a repetitive, sustained, stereotyped pattern. Must travel the same route at least 3 times. Includes travel patterns interrupted by other repetitive behaviors, e.g., if the animal stops and looks out of his pen in the same location as part of the travel pattern.
3b	S	Quasi-Stereotypic Pacing	As in 3 above, except animal need not take the same path 3 or more times in a row. Any pacing in which a predictable pattern emerges. There may be variations in the routine or the animal may alternate between a limited number of travel paths. A good rule of thumb to use is to ask yourself whether the animal would wear a path if grass were growing there.

4	S	Climb (substrate)	Vertical ascent or descent, as into trees or structures; counted as climb only if all four extremities leave the ground. Includes only active climbing; if an animal rests in a tree, for example, “rest” is recorded.
5		Stereotypy: (Evolving definition)	Animal engages in unvarying and repetitive acts that have no obvious goal or function. As these can be highly individualistic, new stereotypies will need to be added to the ethogram as stereotypic patterns are discerned for each animal. Context: Record any external stimuli which appear to evoke stereotypy bouts. (Record the occurrence of these stimuli even when they do not evoke bouts of stereotypy.) * Be on the look out for “loose stereotypies.” i.e., those that contain fixed, predictable components with variable, unpredictable elements (Mason, 1995) and repetitive patterns that occur over longer time scales. Since stereotypies develop from normal behavioral elements, the occasional occurrence of one of the following behaviors should not be scored as a stereotypy unless it becomes part of a rigid, repetitive routine.
5a	E	Pirouette	Stands on hind legs and spins at least 90 degrees.
5b	E	Head-toss	Animal abruptly lifts head upward and/or to the side in a swinging movement; often occurs during pacing (especially when turning). Count each individual head-toss.
5c	SLE	Bizarre limb movements	? Not observed in Wolong, but be on the lookout for them.
5d	SLE	Self-biting	Animal bites itself repeatedly in a stereotyped manner (not grooming).
5e	E	Truncated somersault	Places head to ground as if to do a somersault, but does not complete action; often part of a stereotyped locomotor routine.
5f	S	Stereotyped masturbation	Animal masturbates repetitively and rigidly for long periods; may be manual or oral.
5g	SLE	Weave	Animal “weaves” back and forth, undulating the front portion of the body, somewhat reminiscent of a fish’s swimming movements.
5h	SLE	Sway	Animal swings head back and forth, but does not have undulating movements associated with swaying.
5i	1-0 & at beep	Tongue flick	Animal sticks tongue out repetitivelyin a licking movement; not associated with feeding or grooming after feeding.
5k	E	Sit-up	Animal lies on back, then sits upright. May repeat more than once. This behavior mustbe part of a (quasi)stereotypic pacing pattern.
5m	S	Paw suck	Repetitive, sustained sucking of the toes/paw--not associated with grooming. (As seen in Zhuang Zhuang)

5o	SLE	Cage Climb	Animal stands bipedally and sways or makes climbing motions, as if attempting to escape.
5p	S	Stereotypic Regurgitation	Animal vomits and reingests vomit repeatedly (usually 2-4 times/min). Only score for habitual regurgitators, not the occasional bout of illness.
6		Maintenance:	Comfort behaviors.
6a	S	Scratch	Scratch self with paws.
6b	S	Scratch against surface	Animal rubs a small part of its body repetitively back and forth against an object, as if “to scratch an itch.”
6c	S	“Dirt” bathe	Animal rolls on ground, scratching body on surface; may use paws to throw dirt over body. If dirt bathe on substrate other than dirt, score 6c and identify substrate. If dirt bathes on enrichment substrate (e.g.hay), preface with an “E.” Differentiate from locomotor play (rolling), where the point does not seem to be to rubthe body against the ground, and from rolling, which is an estrous behavior accompanied by bleating and/or chirping, etc.
6d	S	Licks vulva	Licks vulva; includes dark region surrounding vulva. If female is turned away so that a clear view is not available and appears to be licking in the general vicinity of the vulva, score 6d (unless she has an infant that may be located in the vicinity of the vulva). Look for wiggling of the ears to determine if she is licking.
6f	S	Licks nipple	Licks nipple
6g	S	Licks/bites	Licks or bites self to clean or maintain pelage. Simply licking the lips is not scored as “6g” unless it is incorporated into a bout of fur licking.
6h	S	Licks/bites forepaws	Licks or bites forepaws in a non-stereotypic, non-play context.
TEX	1-0 and at beep	Tactile Exploration	The use of front or back paws to manipulate a non mobile object such as a wall, log or tree trunk. The focal animal must be stationary, and uses paws to bat, rub or paw at substrate. Do not confuse with scratching wood (30).
OEX	1-0 and at beep	Oral Exploration	The use of the mouth to manipulate a non mobile object such as a wall, log or tree trunk. The focal animal must be stationary, and uses mouth to bite or chew at substrate. Score only for cubs; for adults, use 29 (chewing). Do not score OEX if the animal is using its mouth to aid in climbing/attempting to climb.

7a	at beep	Locomotor play (at beep)	Superfluous, apparently purposeless activity such as, gamboling, frisking, somersaulting, rolling, and leaping. Record at point samples only. In addition, record the behaviors listed below using 1-0 sampling per 1 min interval.
PR	1-0	Playful running	Rapid playful locomotion (gamboling, frisking)
JU	1-0	Jump	Jumping, rearing up, or bucking in play; may twist and turn body.
SS	1-0	Somersault	Rolls head first, moving forwards; must complete roll.
HS	1-0	Head Stand	Stands on head; hind feet may be off the ground with body extended or body may be curled with hind feet on ground.
RO	1-0	Roll	Rolls and/or twists from side to side. Must turn over at least 90 degrees.
SB	1-0	Self Bite	Panda bites self in a playful, not stereotypiccontext, i.e., it should be accompanied by other play behaviors, such as rolling, head stand, or somersault.
HA	1-0	Hang/Dangle	Panda climbs vertically (e.g., on tree, cage bars, etc.) and hangs/dangles by one or more paws; at least two paws must be free. Animal is not stationary, but wriggles or swings about.
INO	1-0 and at beep	Investigates Object	Panda investigates object by placing nose close (< 3 inches) to object and sniffing and/or licking. Note that INO differs from 12 in that it is scored using 1-0 method. Score E-INO, not E-FS, when panda is investigating a food enrichment item (e.g. fruitsicle, puzzle feeder). INO takes precedence over 7b.
7b	at beep	Object Play / Interaction	Dragging, batting, tossing or manipulating objects. Record at point samples only. In addition, record the behaviors listed below using 1-0 sampling per 1 min interval. None of these behaviors are mutually exclusive, and several are likely to occur in a given play bout. Always record object type (twig, rock, etc.). Preface each entry with an E if interacting with an enrichment item. If a cub is interacting with bamboo, include the suffix “-bam” with any all-occurrence and point sample behaviors (ie, MAN-bam, 7b-bam).
MAN	1-0	Manipulates Object	The panda picks up, holdsand manipulates object by rotating it, pawing at it, or any other play-like behavior which does not fit into other categories. This includes lifting a portion of a large object (e.g. one end of the spin feeder) off the ground or otherwise supporting weight (as with a hanging object).

PAW	1-0	Paws Object	Uses paw to push, swat, or roll an item on the ground, or hangingsuch as boomer ball, PVC rattle, or burlap doll. This can also be associated with food searching, if the panda pushes a puzzle feeder, fruitsicle or feeder ball while trying to get food out.
HPO	1-0	Head Push Object	Panda pushes object with head/muzzle, causing it to move.
BO	1-0	Bite Object	Bite or mouth object. Includes vigorous biting and gentle gnawing (note that this replaces the old “29”, i.e.., chewing).
TO	1-0	Throws Object	Uses paw or mouth to throw object at least 0.3 m
ROO	1-0	Rolls With Object	Rolls, wriggles, head stands, or somersaults--as defined above--while holding object in paws or mouth and/or rolling on top of the object or substrate. Note that ROO is context based; if the interaction appears to be scratching, score 6c.
SO	1-0	Shakes Object	Holds object in mouth and shakes head vigorously.
JUO	1-0	Jump on Object	As in JU above, but with object. Jumps/pounces onobject forcefully, perhaps pushing down on the object with forepaws or grabbing at it.
LWO	1-0	Locomotes With Object	Carries or drags object in mouth or paws.
PRO	1-0	Playful Run w/ Object	As in PR above, but carries object in mouth or paws.
PUR	1-0	Pursue	Animal HPO's or PAWs object, causing it to move, and pursues it a distance >2m.
HAO	1-0	Hang/Dangle with Object	As in HA above, but while holding object in mouth or paws.
7c	at beep	Water Play	Splashing and rolling in water. Record at point samples only.
WSP	1-0	Water Splash	Splashes water with forepaws or hindpaws.
WRO	1-0	Water Roll	Rolls in water.

7d	E & at beep	Social Play	Any "playful" interaction: characterized by lack of apparent purpose, exaggerated and vigorous movements, "incompleteness" in comparison with adult behavior, frequently repeated motor patterns, etc. Social play generally should not be accompanied by aggressive vocalization, though aggressive vocalization may terminate a bout of play-- especially if it becomes too rough. In order to score social play at least one of the behaviors listed below must occur. Record who initiates each play bout (i7d), whether the partner responds with social play (r7d), and who terminates (t7d) each play bout. Record at the beginning of a play bout and at point samples. In addition, record the behaviors listed below using 1-0 sampling. Also, indicate the first behavioral action that begins the play bout (e.g., i7d-PS or i7d-SRO). If you don't know what started it, don't guess! Who does what? It will be too difficult to score what each play partner does (except when using videotape), so just record the focal's behavior-- note that many behaviors imply what the social partner is doing anyhow (e.g., SRO, MF). There are two exceptions: 1. Always record all play behavior by both the mother and her infant. 2. Always record who initiates play AND whether the partner responds with play. Example: Tian Tian is focal, Ling Ling is partner. TT-i7d-PS, TT-i-RU, LL-r7d, TA, SRO, NBH, HS, CH, LL-t-7d. Note that all behaviors not prefaced by a panda's name are assumed to be the focal's actions. Note that play takes precedence over other behaviors, e.g., 2.
i7d	E	Initiates Social Play	Always record who initiates social play.
r7d	E	Responds w/ Social Play	Record whether the other pandas responds with social play.
t7d	E	Terminates Social Play	Record who terminates social play.
CLP	1-0	Climb Panda	Climbs on top of panda (at least 3 paws on panda).
SRO	1-0	Social Roll	Partners engaged in rough and tumble play in which two animals are rolling together such that their relative positions change, i.e., the one on the bottom rolls to the top and vice-versa.
RI	1-0	Roll Invitation	Animal initiates play by rolling on its back to encourage partner to began a play bout.
RU	1-0	Rear Up	Animal rears up on its hind legs, facing, and in close proximity or touching another panda.
TA	1-0	Tackle/Push	Animal tackles or attempts to tackle or knock the opponent off balance by pushing against the opponent with the forelimbs, head or shoulders.

HB	1-0	Head Butt	Animal "walks into the other bear with its forehead and gives a quick head lift movement trying to knock the other bear off balance" (Henry & Herrero, 1974).
PS	1-0	Paw Swat	Animal bats the other with paws. Actual physical contact of individual swats is brief.
BT	1-0	Bite	Animal bites the other. Context is playful: minimal, if any, pain is inflicted.
NBH	1-0	Neck-Bite-Hold	Animal grabs the other by the neck and holds on for > 3 seconds.
SH	1-0	Shake Head	While NBHor BTthe animals shakes its head vigorously, pulling the opponent's skin.
TWR	1-0	Twists and Rolls	Animal twists and rolls in an attempt to break the NBH.
HLC	1-0	Hind-Leg Claw	To escapeNBH, the animal "lifts its hind feet and alternately claws at the forehead of the opponent" (Henry & Herrero, 1974).
MF	1-0	Mouth Fencing	Opponents grab each other by the mouth, interlocking their jaws, or attempt to get a hold of each other's muzzle, perhaps using "fencing" movements.
CH	1-0	Chase	Vigorous pursuit of another individual for > 3 adult panda body lengths. Context is playful.
FL	1-0	Flee	As in CHabove, except that focal is being chased.
		Play Signals????	Be on the lookout for potential play invitation signals!! Here's a few to look for: --play face --play vocalization --ear position --gambol -- roll over on back --rear, paw swat, play bite, head butt, lunge (bears) -- bouncy approach -- head toss, play bow, somersault
7f	S	Sexual Play	Sexual behavior by young animals in a playful context, as suggested by lack of sexual vocalizations and incomplete versions of adult sexual behavior. Sequence may differ from adult behavior.
MT	1-0	Mount	Mounts in which the genital areas are brought into contact or near contact. May grab the partner's scruff in the mouth and grasp the sides with the forepaws. DON’T CONFUSE THIS WITH CLP!
PT	1-0	Pelvic Thrust	Thrusts pelvis (as in copulation) while mounted as defined above.
VS	1-0	Vulvar Stimulation	Gentle licking and biting of the genital region of the female. Not to be scored by mother's licking the genitals of her offspring.

8	S	Rest	Lying or sitting down, either awake or asleep.
9	SLE	Anogenital mark (substrate)	Rubs anogenital region (tail up) against object or substrate. Note (1) the substrate (a, b, c, etc.) (2) whether accompanied by urination by placing a “-10” after the scent mark code and (3) for 9dwhether onlyurination takes place, i.e., little or no contact is made between the anogenital gland and substrate. For contact, record "C" for no contact record "NC." ex: 9d-c-10-NC (8s) *Record duration of all scent marks!
9a	SLE	Squat	In squatting posture. Seen in males and females, but more common in females.
9b	SLE	Leg cock	With one hind leg raised. I have never seen scratching in this position.
9c	SLE	Reverse	Backs into a vertical surface before marking. This scent mark can be confused with scratching, especially for Shi Shi. Scent marks generally have more of a circular motion whereas scratching tends to be back and forth or up and down. Also, scratching bouts tend to last much longer than scent marks, which rarely exceed 15 seconds in length. The crucial component is that the scent gland clearly makes more than just incidental contact with the surface. Seen in males and females, but more common in males.
9d	SLE	Hand stand	Elevates hind quarters vertically, hands support weight; both hind feet must leave the ground. Note whether anogenital contact is made(NC=No Contact, C=Contact). Only seen in males.
10	SLE	Urinate	Passing of urine.
10-3L	SLE	3-legged urination	Stands in quadripedal position, but lifts one leg off the ground while urinating. Often bounces while urinating, reminiscent of urination in 9d posture.
11	SLE	Body rub (substrate)	Other than anogenital mark, rubs areas of body (head, neck, chest) against structures or substrate, in a smooth, fluid motion in which the animal rubs a large portion of his back, especially the neck and shoulders and sometimes the tail, against an object in a way that seems to be intended to impart scent onto the object (usually seen only in males). Often associated with social interaction or olfactory investigation. When body rubbing with the tail, distinguish from scratch and scent-mark (in which tail is up).

12	SLE	Olfactory investigation (substrate)	Places nose close to the substrate and sniffs and/or appears attentive for > 1 sec. Sniffing of the air is not included in this definition. If the animal is at cage bars, it is sometimes difficult to distinguish between 12d and 17a. In 17a the animal moves its head slightly upward in sniffing movements and does not hold its nose to the surface. The animal also appears more attentive to what lies beyond the bars. Note substrate. Note if the following occurs during the OI bout: -fle= “flehmen” = raises lips, exposing teeth and/or opens mouth while investigating scent. Score flehmens in a "one/zero" manner per minute. Regardless of how many times they flehmen in one minute, score it only once. -lick= licks area that it is sniffing -flehmen and lickin the same bout * Please be conservative when scoring "flehmen" and "lick." Any apparent flehmen or licks associated with feeding should not be scored. "Flehmen" usual only occurs when an animal is investigating another's scent. The animal opens it's mouth and sucks air into it's oral cavity, often drools prolifically, and is intently focused on the scent. Please review the training video. * a "12" CANNOT be scored in a one-minute interval BEFORE, DURING, or AFTERa "1b" is scored. (see protocol for an more in-depth discussion on scoring OIs and food directed behavior)
SA	SLE	Scent Anoint	Picks up object w/ paws and rubs object over body. (Note difference between SA and body rub; also note that dirt is notconsidered an object).
14	SLE	Defecate	Passing of fecal matter.
15	E	Startle	Sudden, intense movement such as whole body jerk, indicative of surprise or fear. This is not a response to another panda but to an inanimate object or sound. *Describe disturbance or precipitating event which caused response.
16	SLE	Withdraw	Any form of attempted distancing from an outside stimulus (not another panda), e.g., rapid retreat, cringe, hide. *Describe disturbance or precipitating event which caused response.
17a	S	Stationary alert	Alert, standing quadrupedally, sitting or lying quietly, but remaining attentive, moving head from side to side and/or sniffing air, perhaps attending to external stimuli. If, for example, a locomoting animal pauses and sits quietly for a few seconds, it would almost always meet the criteria for “17a.” Simply opening the eyes and/or shifting position while lying/resting is not sufficient to merit “17a” designation. If, while lying, the animal raises its head and appears attentive to its surroundings, this should be scored as “17a.” Note that this behavior lies on a continuum with rest.

17c	S	Water contact	Sits, stands or lies quietly in water; no water play. Must have at least one paw in the water. This state also applies when panda is sitting or standing with at least one paw in the drinker(for at least 5 secs.).
17d	SLE	Bipedal (substrate/ context)	Stands on hind feet; often accompanied by sniffing movements (directed to air, not substrate) and exploratory behavior. Panda's entire rear end must be off the ground.
17e	S	Door-directed	Panda at the door/gate, behavior oriented toward food, keeper, or bedroom area. Supersedes any behaviors that could be embedded in door directed behavior, i.e. 17a, 6a, 12d, 17d, etc. Simply sleeping or feeding by the door is not sufficient to score a 17e. To be scored in the EXHIBIT, the above behavior must occur within one body length of the door. For the YARD,the above behavior must occur within two body lengths along the fence.
17e-1		Door-directed Mild	Panda waits at the door with mild restlessness. This includes investigating the door, stationary alert at the door, scratching himself, or looking on the other side of the door.
17e-2		Door-directed High	Panda is door directed but with a high degree of restlessness. This would include pacing back and forth in front of the door, pushing and manipulating the door, shifting posture frequently while waiting, vocalizing frequently, or other behaviors that indicate a high anticipatory level of food, keepers, etc.
18	S	Drink	Takes in water.

20		Vocalization: (context)	SCORE USING 0NE-ZERO SAMPLING METHOD. Record “20” for unidentifiable vocalizations, in parentheses make your best guess as to call type on the ethogram. Only record the vocalizations in bold below. The other vocalizations will only be scored as part of transactional behaviors.
20a	1-0	Bleat	A twittering, goat-like call of variable length (1-3s). Contact call, appeasing, non-aggressive, “friendly.”
20b	1-0	Chirp (yip) OR Sex Squeal	Short, tonal, high-pitched, descending in pitch toward end. Context 1: Most common during peak estrus. Friendly, promotes social proximity. Males will rarely chirp, usually when in a state of heightened excitement when first interacting with an estrous female that has just been relocated to the pen next to them. Do not confuse with a truncated bleat. Context 2: Several subadult males in Wolong will chirp, or more commonly give a much louder version called a "yip," usually when they are highly aroused about being confined in a small space and/or when anticipating feeding time. This vocalization (yip) is a piercing call that hurts your ears if you are very close. Contextually, it resembles honking, except that the animal appears much more agitated. Similar to agonistic squeal, but is more harmonic. The context is key in differentiating between the two, the sex squeal is affiliative where as the agonistic squeal is antagonistic. Gradations between chirps and squeals sometimes make these vocalizations very difficult to distinguish, thus both are lumped under "20b." Also, a similar vocalization is heard by mothers with infants.
20c	1-0	Bark	Short, (0.1-0.3s), fairly noisy, similar to dog bark. Threat, causes withdraw by receiving animal.
20d	1-0	Moan	Low-pitched, low-to-medium amplitude, call of variable duration. Often has several short starting elements. Often grades with growls, barks, chirps and bleats. “Ranges from a soft hoot and softly repeated bu-bu-bu to a low-pitched moo, whiny groan and long drawn-out moan rising and falling in pitch” (Schaller et al, 1985). Mild threat, often used by the female to discourage approach by the male. Perhaps best described as a “wookie call” as exemplified by Chewbacca of Star Wars fame.
20e	1-0	Growl	Long, noisy, low pitched growl similar to a dog’s. Aggressive, often accompanies or precedes fighting. Only used by attacking animal.
20f	1-0	Honk	Short (< 0.5s), tonal, low pitched, nasal, falling in pitch, produced repetitively in a series. Often used when stressed or frustrated.

20g	1-0	Roar	Very loud, intense, harsh (highest level threat).
20h	1-0	Chomp	Alternate rapid opening and closing mouth, with teeth coming hard together. Defensive, e.g., when the female avoids the male’s approach. Mild threat.
20i	1-0	Huff	Audible expulsion of air through open mouth (anxious, mild threat).
20j	1-0	Snort	More intense expulsion of air through nose (threat, apprehension).
20k	1-0	Agonistic Squeal	Short, high-pitched, open-mouthed call. Often used by subordinate animal in a fight or in response to pain.
21	1-0 & beep	Tail Up	Tail raised to uncover vaginal opening. Often accompanied by bleating and searching, used by females in estrus to indicate receptivity to mating (part of lordosis posture) If tail up is caused by a person, record "31 (tail up)." Score using one-zero sampling method and at beep!
22	E	Foot scrape	Rapid scraping of hind feet back and forth on substrate while standing in place; aggressive act.
23	S	Genital Manipulation	Any rubbing or grasping of genitalia with paws. Note: score “23” for manipulation of vulva by female prior to birth event.
24	SLE	Backwards walking	Female walks backward for several steps. Usually seen during estrus. Often toward male and may be followed by anogenital present. Often accompanied by bleating and chirping. Note if backwards walking occurs through water and record as in '2-Locomote' above (i.e. 24-g).
26	1-0 & beep	Lordosis	As in 21, but hind quarters raised, and kneeling on forelegs; head often on the ground. Score using one-zero sampling method and at beep!
27	SLE	Rolling	Animal lies on back and rolls and writhes (as if back is itching). Common in females reaching peak estrus.
28	SLE	Digging	Animal digs in dirt or loose substrate with forepaws.
29	SLE	Chewing	Animal chews on wood or other non mobile substrate, such as logs, trees, and doors.
31	S	Human Oriented	Approach and observe person closely; may stand bipedally or interact with human in any way. Does not include simply looking toward the visitors.

32		Other	Any behavior which does not fit definitions on the ethogram, for example, olfactory investigation of food. Describe.
FOL	S	Follow	A purposeful, directed pursuit of another panda for at least 3 adult body lengths. The animals need not start in close proximity, but should be in visual contact when the “leader” moves away. The “follower” must end up no more than 2 adult body lengths away from the “leader”.

TRANSACTIONAL SOCIAL BEHAVIORS (to be recorded for interactions between adults)

Rule of thumb: An interaction must last at least 5 seconds to be scored.

TE	SLE	Territorial	Panda footscrapes while interacting with another panda. TEaccompanied by any vocalization should be scored according to the appropriate category for that vocalization (e.g., N2), NOT TE.
N1	SLE	Non- Contact Aggression, Level 1	Agitated Interaction which includes mildthreats which are primarily defensive, connoting alarm, distress, apprehension, aversion. Vocalizations include huff, snort, chomp, and honk.
N2	SLE	Non- Contact Aggression, Level 2	Threatening. Interaction which includes moderate threats. Actions include paw swats without contact andcharge/lunge. Vocalizations include moan, bark, growl androar.
C1	SLE	Contact Aggression, Level 1	A non-sustained (i.e., less than 5 seconds) aggressive interaction of moderate intensity, including actions such as paw swat, bite, grab, and dominance mount. “Moderate intensity” is defined as aggressive acts which have low potential for inflicting serious injury, e.g., drawing blood. Generally accompanied by vocalizations discussed in N2.
C2	SLE	Contact Aggression, Level 2	Sustained aggression,includingvigorous and potentially injurious physical contact (fighting, attacking), such as bite, grab, paw swat, wrestle, chasewith contact. Intensity of aggression is high, as indicated by injury, duration, or vocalizations such as roarand squeal.
SI	SLE	Shows Interest	Animal appears "eager" to interact with the other panda, as suggested by pushing and pulling at fence separating the pandas (during PC), sniffing at the other panda, greeting, pacing back and forth in close proximity to the other panda, or circling the other panda with attention clearly focused on the other panda. Score SI when no vocalization has occurred and behavior is clearly not neutral (for example: Shi Shi is eating biscuits; Bai Yun approaches him from behind and sniffs him without vocalizing). SI accompanied by any vocalization should be scored according to the appropriate category for that vocalization (e.g., AF), NOT SI!
AF	SLE	Affiliative	Animal approaches or attempts to interact in a “friendly” manner, as evinced by vocalizations such as bleat, sex squeal, chirp.

SX	SLE	Sexual	Animal approaches or attempts to interact in a proceptive or receptive manner, i.e., attempting to engage in courtship or sexual interaction. Activities include rear present (RP), anogenital present (25), and tail up (21)for the female; and sexual mount, anogenital inspection (AI), and gentle nudging and pawing the female's anogenital area by the male. Generally accompanied by one or more of the vocalizations described in AF. Note that vocalizations alone cannot be used to distinguish Sexual from Affiliative transactions, but must be interpreted in the light of the context and other behavioral acts. SX takes priority over other transactional behaviors, e.g. AF and AM. For example, if a male bleats while anogenital investigating, score SX only, not AF, SX. • Note that if the female begins to masturbate, this terminates the SX interaction. The female is scored as "23." If the male remains in proximity, he is scored as AF, SI, IG, etc. depending on his behavior and the female is scored as IG in addition to 23. 23 should be scored at the beep.
IG	SLE	Ignore/Neutral	Panda shows no overt response to proximity or behavior of another panda. Should be scored when in proximity lasts more than 5 secs. and none of the behaviors described in other categories occur. For example, one animal approaches the other and/or attempts to interact, but the recipient shows no overt reaction or simply observesthe initiator. If focalIG at beep, score the individual behavior of the focal(e.g., 1a).
AM	SLE	Ambivalent	Behavior during transaction includes both friendly and aggressive overtures, i.e., includes elements of N1, N2,or C1 and AF or SX. For example, the female may approach and moan, bark, chirp and bleat. In parentheses, please record the combination of aggressive and friendly categories, e.g., AM (N2 and AF)
AV	E	Avoid	Behavior intended to prevent or avoid interaction. Panda turns away, backs off, or redirects its travel path to avoid interacting with another panda in proximity (< 2 BL). Behavior serves to increase inter-animal distance or avoid close proximity. For example, a panda whose travel path is approaching another panda will often abruptly change its direction of travel, usually accompanied by sidelong glances and hesitation in gait at the point of travel path alteration. Simply pacing back and forth in front of another panda does not constitute an avoid; instead it should be scored SI. Note that a female turning around to present her hindquarters to the male (anogenital present) would notconstitute an Avoid
RT	E	Retreat	Same as AV above, except panda is “forced”to increase the distance between itself and the other panda, i.e., in response to aggressive activity by the opposing party, such as that discussed in N1, N2, N3, C1, C2, AM.

SUB	SLE	Submission	Animal adopts a submissive posture (facing away from the other panda, crouched with head down) or simple turns whole body away from the other panda, not increasing social distance. Animal may or may not submit in the face of attack(C1 or C2) or threatening vocalizations by opposing animal. SUB accompanied by any vocalization should be scored according to the appropriate category for that vocalization (e.g., N2), SUB!
AP	E	Approach	Panda moves to < 2 body lengths from other panda. An approach alwaysbegins an interaction sequence
DP	E	Depart	Panda moves to > 2 body lengths from other panda. A depart alwaysends an interaction sequence. Note: if panda "departs" for less than 10 seconds and then "re-approaches" the other panda, a depart is not scored and the interaction continues.

The following behaviors were used in an earlier ethogram. Here these behaviors are defined simply for reference to clarify definitions of transactional behaviorsdescribed above. There is no need to record these behaviors.

		Observe	In exhibit (PC or DC): Approaches cage door separating adjacent pen and looks into neighboring enclosure, regardless of whether occupant is visible or not. In exercise yard or Direct Contact: Animal directs gaze toward other animal, which is in its field of view.
		Spar	Joint directed activity without sustained physical contact. The animals might be sitting or standing facing each other or standing on their hind legs and paw swatting at each other or periodically attempting to bite (NZP, 1992). Score bites and paw swats as events embedded in SP state. Don’t worry too much about when one bout begins and ends, because it’s the point samples that count most. Ex: SP-ps-ps-bt-ps-bt (remember, 5 sec. rule applies for separating bouts).
		Wrestle	This activity includes sustainedcontact and is usually more intensive than sparring. One animal is often mounted on top of the other, or they may jointly wrestle or grapple with each other (NZP, 1992).
		Aggression	Vigorous wrestling, striking and biting which could potentially do bodily harm. Includes attempts to engage in such activity through cage door, and such interactions accompanied by barking, growling and squealing.
		Gree	Naso-nasal contact or near contact, often accompanied by mutual sniffing.

		Paw swat	Animal bats the other with paws. Actual physical contact of individual swats is brief. Note that for behavioral events, boutsmust be separated by at least 5 seconds before scoring a second time-- see protocol.
		Charge/Lunge	Rapid aggressive approach of another individual or rapidly turning to face a nearby animal, decreasing the distance between the two animals.
		grab	Animal grabs, claws, or holds other animal down with forepaws.
		Bite	Animal bites the other. May occur in aggressive or playful context.
		Chase	Vigorous pursuit of another individual for > 5 panda body lengths. Context is aggressive.
		submissive posture	panda sits with hind quarters oriented toward other participant in a crouched position following a social interaction. Generally, following an agonistic interaction, the animal will turn around to avoid further confrontation
		dominance mount	In an agonisticcontext, panda rears up and places paws on the hind quarters or back of other participant, often pushing down or pushing away. Context is aggressive and should be accompanied, preceded or followed by threatening vocalizations or aggressive acts, such as paw swat, charge, or bite. Note that sexual mounts are often accompanied by aggressive behavior. The key to distinguishing dominance mounts is the complete lack of sexual motivation on the part of the mounter(the mounted animal may or may not be sexually motivated)
		Anogenital present	Female presents hind quarters to male (during estrous) with tail up (may or may not be in lordosis posture).
		Rear present (Anogenital present w/out tail up)	Female presents hind quarters to male (during estrous) with out lifting her tail.

Substrates:		Used for climb, anogenital mark, olfactory investigation, body rub, bipedal stance, and point samples. If at the point sample the animal is on two or more substrates, record the substrate which is in contact with (or underneath if the animal is standing) the largest portion of the animal’s body.
-a	Dirt/grass		-g	Water
-b	Cement floor		-h	Stone/Drinker/Cave/ tree guard
-c	Wall		-i	Feces
-d	Cage bars		-E	Enrichment item
-e	Live tree or bush		-k	Hay
-f	Dead wood		-P	Panda

Location:

Used for point in time samples. If at the point sample the animal is in two or more locations, record the location in which the largest portion of the animal’s body is contained.

DOOR/GATE: If panda is within one body length of door or gate then at the beep, record the location as "d/g". In the EXERCISE YARD,the door/gate is defined as the gate itself plus 2 panda body lengths along fence.

INSIDE:

P	Platform	Raised sleeping platforms in bedrooms
B	Bedroom
S	Sun Room	Small outdoor enclosures adjacent to bedroom
G	Garden Room	Larger enclosures with grass; adjacent to sun room

ENCLOSURE: Used when experimental condition is "CHOICE" only.

Y	Exercise yard
E	Exhibit
T	Tunnel
B	Bedroom
S	Sun Room
G	Garden Room

BIRTH WATCH / MOTHER-INFANT INSTRUCTIONS

OBSERVATIONS:

Observers for birth watch sessions should record behavioral data in the same manner that observations are usually made and using the new birth watch datasheets. Following are a few changes which need to be carefully followed

• In contrast with our behavioral observations on adults, we will score all occurrences of bouts of vocalizations for infants

•We will also score infant vocalizations at the beep

• This is a new ethogram, so added descriptions are even more important; pay attention to details.

• After the birth, always preface mother behavior with an “M-” and infant behavior with an “I-”

• If a behavior by the infant immediately precedes a response by the mother draw an arrow between the infant’s behavior and the mother’s. Likewise if the mother’s behavior causes a change in the infant’s behavior draw an arrow linking the two. Pay close attention to the order: who changed their behavior first, the mother or the infant?

Example:

Interval 1:(1) Mother and infant are resting quietly; (2) infant begins to squawk and you notice that the infant has become partially visible, exposed to the cold; (3) the mother shifts her position, places her nose close to the infant, and covers the infant with her upper arm; (4) both animals rest quietly, then the watch beeps;

Interval 2: (5) the mother changes her position, rotating 180°, and then the infant cries for 15 seconds; the mother gently cuddles the infant and carefully covers it up with her upper arm; (6) mother and infant rest quietly and the watch beeps:

Interval 3: (7) the infant begins squawking; (8) you can’t see the infant, but the mother clearly attends to the infant as evidenced by shifting her posture and placing her nose closer to the infant; (8) the watch beeps.

Interval 4: (9) The mother rotates about 60° and (10) shifts infant from her paw to under her chin. You can no longer see her face nor the infant, but you know where on her body the infant is located. (11) You can tell from the wiggling of her ears that she is licking, but you can’t tell where. (12) Beep.

Min	All Occurrences	M-Bhvr	I-Bhvr	I- Voc	Contact	Exposure	Location
1	M-8, I-8, I-20p* ->M-CO	8	8	NO	HH	NE	Den
2	M-CP ->I-20m ->M-CO, M-8, I-8	8	8	NO	HH	PE

Behavior Ethogram (San Diego Zoo) and Protocol