behavior of female african elephants (loxodonta africana) in captivity
TRANSCRIPT
Applied Animal Ethology, 6 (1980) 257-276 257 0 Elsevier Scientific Publishing Company, Amsterdam - Printed in The Netherlands
BEHAVIOR OF FEMALE AFRICAN ELEPHANTS (LOXODONTA AFRICANA) IN CAPTIVITY
JACK ADAMS and JUDITH K. BERG
California State Uniuersity, Dominguez Hills, Carson, CA 90747 (U.S.A.)
(Accepted for publication 26 June 1979)
ABSTRACT
Adams, J. and Berg, J.K., 1980. Behavior of female African elephants (Loxodonta afri- cana) in captivity. Appl. Anim. Eth@., 6: 257-276.
The purpose of this research was to ascertain the repertoire of behavior of female African elephants (Loxodonta africana) in captivity. Seven female African elephants were observed for 558 hours and 22 minutes for a period of one year. This paper gives a detailed description of the activities of the elephants maintained in a relatively re- stricted environment. Twenty one different kinds of behaviors were observed, 14 of which were considered unique to elephants. The most frequently occurring behavior was the placement of the trunk of one elephant into or near the mouth of another elephant. The activities were discussed in terms of: (1) social behaviors; (2) individual behaviors; (3) biological behaviors; (4) dominance hierarchy; (5) four factors derived by statistical factor analysis.
INTRODUCTION
African elephants have been kept in captivity for a long time. The earliest account described them in use in war by the armies of Hannibal in 218 B.C. Since then they have been used in ceremonial pageantry, for work in logging and transportation operations, for entertainment by circuses, and on display in zoos. However, there is relatively little scientific information regarding their repertoire of behavior in captivity. This is probably attributable to their massive weight and enormous size. The biggest African elephant in captivity was “Jumbo” owned by a famous American circus. It weighed 12 000 lbs and its height was 10 feet, 9% inches. In addition, elephants have been known to attack and physically injure, and sometimes even kill, their caretakers. They have also demolished large structures when on a rampage. Therefore, elephants were considered to be potentially dangerous animals. Consequently, they were kept in relatively safe but small enclosures which facilitated their management and husbandry. Such enclosures did not pro- vide sufficient opportunity for elephants to demonstrate a wide range of behaviors. A review of the literature revealed that the care and management
258
of orphan elephants have been reported by Reuther (1969), Young and Oelofse (1969) and Sheldrick (1975). Limited reports on the behavior of African elephants were found in publications of Rensch (1957), Kuhme (1962), Hediger (1964), Bolwig (1965), Bolwig et al. (1965), Hediger (1968) and Adams and Berg (1974).
With the advent of “Wild Animal Parks”, where elephants are given relatively more freedom to move around in significantly larger open areas under con- trolled conditions, it has become feasible to observe and quantify the variety of their daily activities.
Five of the seven elephants reported on in this study were of special inte- rest because one of us (JA) witnessed and filmed their capture and taming procedure at Wankie National Park in Rhodesia in 1971, before they were shipped to their present location (Adams, 1973).
The purpose of this study was to ascertain the repertoire of behavior of female African elephants in captivity at the San Diego Wild Animal Park.
MATERIALS AND METHODS
Subjects
The subjects were seven female African elephants located at the San Diego Wild Animal Park, San Pasqual, California. They were identified as follows:
Name Age
Hatari Sabu Deteema Madundamella Mandavu Shapie Wankie
15 years (young adult) 14 years (young adult)
5 years (calf) 5 years (calf) 5 years (calf) 5 years (calf) 5 years (calf)
The elephants could reliably be distinguished by their appearance.
Apparatus
All the elephants were contained in a moat-surrounded area, 625 feet (190.6 m) long and 225 feet (68.6 m) wide (Fig. 1). The elephant com- pound was oval in shape and it narrowed to a la-foot (3.6 m) gate at one end. The enclosed area contained two large concrete shade structures and a concrete water “hole” containing approximately 65 000 gallons (245 025 1) of water.
The elephants were kept chained in a barn adjacent to the described en- closure during the night hours. They were released into the enclosure each day at approximately 08.30 h and they were allowed to enter the barn at approximately 17.00 h daily.
Fig. 1. Portion of the moat-surrounded African elephant enclosure at the San Diego Wild Animal Park, showing a concrete structure for shade and security, and tree stumps for rubbing.
Procedure
Observations of the elephants were made from a point outside the en- closure giving an unrestricted view of the whole area. Tape-recordings, still- and tine-photography was used when appropriate.
The period of observation was one year, commencing on 23 May 19’75 and ending on 22 May 1976. There were 24 sessions of observations of 3-4 days duration, on 85 different days, for a total of 558 h and 22 min. The shortest period of observation was 2 h 30 min, and the longest was 9 h 35 min. The average amount of time spent in observing the elephants was 6 h 30 min.
RESULTS
Twenty one separate activities were observed in the repertoire of behavior of the female African elephants. Table I shows the rank order of the fre- quency of occurrence of 14 behaviors which are considered unique to these elephants. A summary of the 14 behaviors observed, according to the rank order of frequency of occurrence by each individual elephant, is shown in Table II.
The actual activities comprising the repertoire of behavior are given in detail below.
260
TABLE I
Rank order of frequency of occurrence of specific behaviors of African elephants in captivity in the San Diego Wild Animal Park’ ___~ _~ ~~ - Behavior Frequency Mean ___~ ~~ ___--
(1) Trunk to mouth 1540 220.0 (2) Dusting 1321 188.1 (3) Trunk intertwining 720 102.9 (4) Reaching across moat 624 89.1 (5) Investigation and manipulation 389 55.6 (6) Trunk placement 372 53.1 (7) Eating objects 306 43.7 (8) Lying down 289 41.3 (9) Mudding 242 34.6
(10) Temporal gland secretion 187 26.7 (11) Knocking trunks 101 14.4 (12) Vocalizations 97 13.9 (13) Rubbing 93 13.3 (14) Agonistic 41 5.9 __ ‘See text for eight additional behaviors observed but not tabulated.
(1) Trunk to mouth activity was the most frequently observed behavior. It involved the placement of the trunk of one elephant into or near the mouth of another elephant (Fig. 2). This behavior was observed under the following conditions: when the elephants were eating or chewing on something; when one elephant was drinking; when passing one another; when the elephants approached each other; before or after intertwining trunks; when frightened; when walking side by side. Trunk to mouth behavior was frequently reci- procated. Some elephants engaged in this activity more than others, and it appeared to be selective as to which elephants participated in it. The highest incidence of this behavior was between Sabu and Hatari (the two young adults). Most of this behavior was initiated by Sabu. Except for placing her trunk to the mouth of Wankie (the youngest and smallest calf) for a total of 72 times, most of Sabu’s participation with other elephants in this regard was very minimal, i.e. to Mandavu 8 times, to Deteema 8 times, to Shapie 6 times, and to Madundamella 2 times. It appeared that there was a special relationship between Sabu and Hatari, and between Sabu and Wankie rela- tive to the trunk to mouth behavior. It is important to note that Wankie placed her trunk to the mouth of Sabu 117 times, which was the second highest frequency of this behavior. Although all the elephants engaged in the trunk to mouth behavior, Madundamella and Mandavu never placed their trunk to the mouth of Sabu, and Mandavu never extended her trunk to the mouth of Hatari.
(2) Dusting. The elephants spent a considerable amount of time in dusting their bodies with dry sand. The sand was grasped with the end of the trunk
TAB
LE
II
Ran
k or
der
of f
requ
ency
of
beh
avio
rs p
er i
ndiv
idua
l A
fric
an e
leph
ant
at t
he S
an D
iego
Wild
Ani
mal
Par
k
(1)
f (2
) f
(3)
f (4
) f
(5)
f (6
) f
(7)
f Tr
unk
to
Dus
ting
Tr
unk
Rea
chin
g In
vest
ig.
Trun
k E
atin
g m
outh
in
tert
win
ing
acro
ss m
oat
man
ipul
. pl
acem
ent
obje
cts
Man
davu
31
4 Sa
bu
265
Mad
unda
. 17
4 Sa
bu
224
Sabu
10
4 Sa
bu
103
Shap
ie
63
Wan
kie
298
Hat
ari
229
Shap
ie
118
Man
davu
14
5
Man
davu
57
D
etee
ma
75
Wan
kie
60
Shap
ie
275
Mad
unda
. 20
9 Sa
bu
105
Wan
kie
95
Shap
ie
55
Man
davu
67
D
etee
ma
52
Sabu
22
1 Sh
apie
18
3 D
etee
ma
98
Hat
ari
78
Hat
ari
54
Shap
ie
59
Mad
unda
. 43
D
etee
ma
213
Det
eem
a 18
0 H
atar
i 92
M
adun
da.
61
Mad
unda
. 53
M
adun
da.
25
Man
davu
34
M
adun
da.
134
Man
davu
15
4 W
anki
e 72
Sh
apie
18
W
anki
e 38
W
anki
e 23
Sa
bu
33
Hat
ari
85
Wan
kie
104
Man
davu
61
D
etee
ma
3 D
etee
ma
28
Hat
ari
20
Hat
ari
21
Tota
l 15
40
1324
72
0 62
4 38
9 37
2 30
6 M
ean
220.
0 18
9.1
102.
9 89
.1
55.6
53
.1
43.7
(8)
Lyin
g do
wn
f
Sabu
11
7 Sa
bu
Shap
ie
62
Hat
ari
Hat
ari
46
Det
eem
a M
anda
vu
25
Shap
ie
Wan
kie
20
Wan
kie
Det
eem
a 12
M
anda
vu
Mad
unda
. 7
Mad
unda
.
Tota
l 28
9 24
2 18
7 10
1 M
ean
41.3
34
.6
26.7
14
.4
(9)
Mud
ding
f 41
Sa
bu
50
40
Hat
ari
46
38
Shap
ie
30
37
Man
davu
23
31
M
adun
da.
16
28
Det
eem
a 12
27
W
anki
e 10
(10)
f
Tmp.
gln
d.
secr
etio
n
(11)
Tr
unk
knoc
king
Shap
ie
Sabu
M
adun
da.
Hat
ari
Det
eem
a W
anki
e M
anda
vu
f 42
Sabu
40
H
atar
i 10
W
anki
e 3
Man
davu
3
Shap
ie
2 D
etee
ma
1 M
adun
da.
(12)
V
ocal
iza-
ti
on
f (1
3)
f (1
4)
Rub
bing
A
goni
stic
28
Sabu
30
H
atar
i 13
27
M
adun
da.
15
Sabu
11
15
H
atar
i 13
W
anki
e 6
11
Man
davu
12
M
anda
vu
5 9
Shap
ie
11
Det
eem
a 3
4 W
anki
e 8
Mad
unda
. 2
3 D
etee
ma
4 Sh
apie
1
97
13.9
93
13
.3
f 41 5.9
262
Fig. 2. Elephants engaging in trunk to mouth behavior, the most frequent activity among female African elephants in captivity.
and tossed onto the body over the head, from the right or left side or up the middle onto the chest. This behavior was invariably performed when the elephants emerged from the water and when the temperature was about 80°F or higher. It occurred most frequently between 10.00 and 13.00 h, which was usually the hottest period of the day.
Sabu and Hatari had the highest incidence of dusting. Very often the five younger elephants mimicked the older ones. Dusting was also resorted to when sufficient browse was not brought to the elephants, or when there was a delay in admitting them to the barn for the nightly chain-up and supplemental feeding. Dusting was significantly correlated with temporal gland secretion (r = 0.79; P < 0.05).
(3) Trunk intertwining. This behavior was engaged in mostly by Madunda- mella with Deteema, Wankie, Shapie and Mandavu, in that order. Only Shapie had a trunk intertwining encounter with the older elephant Sabu. AI1 the calf elephants engaged in this behavior with all the other calves, whereas the young adult elephants restricted it to each other, except in the case of Shapie and Sabu.
263
Fig. 3. Reaching across the moat behavior by a young adult elephant with her hind legs located on top of the moat and her forelegs on the concrete side of the moat. Wankie, the smallest calf elephant who was not able to mimick the older elephants, found a way to get into the moat to achieve the purpose of obtaining additional vegetation to eat.
(4) Reaching ucross the moat (Fig. 3). All of the elephants engaged in this behavior, which involved reaching across the moat with the trunk and grasping flowers and any other vegetation and eating it. The smallest ele- phant, Wankie, could not reach across the moat with success. She overcame this disadvantage by finding a way to get down into the moat at a place near the water pond and she walked inside the moat for approximately 40 m to where the other elephants were reaching over. Then she reached up and grasped the vegetation with her trunk and ate it. Reaching across the moat occurred at various times of the day but most often at 11.00 to 12.00 h, and between 15.00 and 16.00 h. These time-periods followed the completion of eating the regular hay food provided for the elephants. Reaching across the moat was significantly correlated with investigative and manipulative behaviors (r = 0.81; P < 0.05).
(5) Investigation and manipulation. The elephants investigated and manipu- lated objects they could touch with their trunk. Sabu was the most prolific in this behavior. Objects investigated or manipulated included tree branches, pieces of wood, hay, the tail of another elephant, rocks, cloth, chain, feces, tree stump, clumps of soil, and the ring in the chain at the cement shade
264
structure. The elephants placed pieces of wood, tree branches, leaves and rocks on their back and head frequently and repeatedly as these objects fell off. Investigative and manipulative behavior was significantly correlated with reaching across the moat (r = 0.81; P < 0.05), with lying down (r = 0.88; P < 0.05), and with temporal gland secretion (r = 0.78; P < 0.05).
(6) Trunk placement. The elephants placed their trunk on to the back of another elephant at the medial aspect of the body from a lateral position and, on to the back in an anterior-posterior plane while standing at the rear of the other elephant. The trunk was also placed in various other positions on the body of another elephant such as the genitalia, neck, head and face, the anus, along the back, across the back, under the ear, the temporal gland, the tail, and also on the elephant’s own tusk. Wankie, the smallest and youngest of ail the calf elephants, received more trunk placements from other elephants than any other elephant.
(7) Eating objects other than food. This behavior included eating sand, soil, small stones and the fecal material (coprophagia), either their own or that of another elephant. This behavior was significantly negatively correlated with agonistic behavior (r = 0.76; P < 0.05).
(8) Lying down. All the elephants lay down at various times of the day, and for different periods of time, ranging from simply lying down and rising immediately to lying for as long as 57 min. No preference was demonstrated for lying down on one side of the body rather than the other, but the least frequent recumbeni position was on the ventral aspect of the body. While they were lying down, the elephants’ eyes were closed and it was presumed they were sleeping. They preferred to lie down on damp soil if available. Hatari would frequently stand beside the body of Sabu or Shapie when they were lying down. Shapie would stand alongside the body of Hatari, Sabu or Wankie while they were lying down. The elephants were never seen all lying down at the same time. The elephants selected sloping ground and lay with their head at the highest level.
(9) Mudding. This behavior included sucking up muddy water into the trunk and then blowing the mixture out over various parts of the body, or tossing damp soil on the body. Although there were no mud holes provided for the elephants, they would make a mud puddle by sucking up water out of the pond with the trunk and letting it spill over the ground. The water and ground were then agitated with the trunk and/or forefoot until mud was produced. The elephants also agitated the water standing in small pools in the elephant enclosure after the sprinkling system was left turned on for ex- tended periods of time, and they blew the mud on to their bodies. Mudding occurred most frequently between 10.00 and 13.00 h, which was usually the hottest time of the day.
265
(10) Temporal gland secretion. The temporal gland is located bilaterally on the elephant’s head between the eye and ear. The rank order of the condi- tions under which the temporal glands appeared to secrete included: unde- termined causes; during “wild” activity and commotion; upon release from the bar,1 in the morning; when a truck arrived with acacia tree branches; when harassed by another elephant; when the elephant trainer entered the enclosure; when loud noises were produced by passing trucks or heavy machinery; after being chained to the concrete structure; and when other elephants produced loud growl and trumpet sounds. The highest incidence of temporal gland secretion was by the young adult Sabu, and the least by the youngest calf Wankie.
We did not observe any significant change in the behavior of an elephant after it placed its trunk on the secreting temporal gland of another elephant.
The temporal gland secretion was significantly correlated with such be- haviors as investigative and manipulative (r = 0.79; P < 0.05), lying down (r = 0.84; P < 0.05), vocalizations (r = 0.84; P < 0.05) and dusting (r = 0.79; P < 0.05).
(11) Knocking trunks. This behavior involved two elephants standing face- to-face a short distance apart and then advancing suddenly until their heads, at the base of the trunk, collided into each other in goat-like fashion. This behavior was selective among the elephants. Certain pairs of elephants en- gaged in it more often than others. Sabu and Shapie appeared to have the highest incidence of this behavior.
Knocking trunks was significantly correlated with lying down behavior (r = 0.80; P < 0.05).
(12) Vocalizations. The elephants produced five different kinds of sounds, namely growls, trumpets, crying out, roars and squeaks. These sounds are identified as vocalizations because they are produced in the throat of the animal. More specifically they are: (a) Growl. A low pitch rumbling sound. It was produced when a truck was driven near the elephant enclosure, when one elephant approached or walked near another elephant, when men were working in the barn and making loud construction noises, when acacia browse was brought into the enclosure, when the elephant caretaker came out of the barn, when Hatari was called by name by the caretaker, when a loud noisy machine was started-up, when the elephants placed their trunk in another’s mouth, in response to another elephant’s trumpet sound, when the caretaker entered the elephant enclosure, and when one elephant layed its trunk across the back of another elephant; (b) Trumpet. This sound is similar to that produced by an experienced and talented human musician on a brass musical instrument called the trumpet, hence its name. There were variations of trumpeting sounds which consisted of mixed air and tone, such as an inexperienced human musician might produce from a trumpet. Trumpeting occurred while the elephants were aggressively striking the
266
steel gates to the barn when acacia browse was not brought to them at the usual time, in response to the trumpeting of Asian elephants in a nearby enclosure, and for no apparent reason; (c) Crying-out. This was a shrieking- ly loud sound. It was produced by Mandavu, Madundamella and Shapie when they were physically attacked by Sabu, the biggest young adult, when the trainer was restraining Mandavu with a rope during a training session, and when a truck was passing near the enclosure; (d) Roar. A prolonged low to high pitch sound. It was produced by Hatari when she entered the barn in the evening, by Wankie and Shapie in response to sounds by Asian elephants nearby, and by Mandavu when the caretaker wheeled a bale of hay into the enclosure; (e) Squeak. A high pitch tone similar to that pro- duced by a chalk scratching a blackboard. This sound was produced by Shapie .
Growls and trumpets were the most frequently produced sounds. The two adult elephants produced more sounds than any of the calves.
Vocalizations were significantly correlated with temporal gland secretion (r = 0.84; P < 0.05) and agonistic behavior (r = 0.94; P < 0.05).
(13) Rubbing. The elephants rubbed their body against the steel gate to the barn, on the concrete structure, and against another elephant’s body. Other areas rubbed with the trunk or with hay or small pieces of wood held in the trunk included the head, eyes and ears. The latter was the most frequently rubbed part of the body.
(14) Agonistic behavior. This category included fighting among the elephants and aggressive acts toward other animals. When the elephants displayed this behavior they whipped the trunk at the intended subject, or they used the tusks to push against a subject. The ears of the aggressor were extended laterally. This behavior occurred when the elephants were eating hay or acacia browse. Hatari frequently walked briskly toward a calf elephant with the apparent intention of being agonistic, but the calf quickly moved away and the behavior was aborted. Hatari chased or pushed Mandavu 11 times. On two occasions Sabu joined Hatari in pushing on Mandavu. During one incident, it appeared that Hatari and Sabu attempted to push Mandavu into the moat. Mandavu cried out and trumpeted as she assumed a kneeling po- sition with the forelegs on the edge of the moat, and she secreted profusely from the temporal glands. The caretaker yelled at Hatari and Sabu by name and the agonistic behavior was discontinued. On another occasion when browse was brought to the elephants, Hatari and Sabu attacked Mandavu. They pushed her against the steel gate with their tusks. Mandavu cried out, trumpeted, and secreted from the temporal glands. On yet another occasion, Mandavu was the last to be released from the barn and into the elephant enclosure. When she entered the compound, Hatari and Sabu attacked her by pushing on her with their tusks. Mandavu cried out, trumpeted and finally escaped. Mandavu was the victim of agonistic behavior on 15 different
occasions. Most often she was eating hay or browse when attacked. Madunda- mella was the object of agonistic behavior 12 times, but to a lesser extent in that her assailants only struck her with their trunks. Hatari, Wankie and Madundamella chased and swung their trunks at birds and ducks that landed in the elephant enclosure.
The elephants also demonstrated violent aggressive behavior by repeatedly backing into the steel gates to the barn. This created considerable noise and most of the elephants then dashed around aimlessly with ears extended, trumpeting, and banging their trunks at piles of elephant droppings or tossing a rock or debris on to their back with their trunk. Sabu engaged in this behavior more often than any other elephant. The striking of the gates, the rapid movements and trumpeting by Sabu stirred up the others and there was considerable commotion, trumpeting and dashing about by all the other elephants in the area. Such behavior occurred most often between 14.00 and 16.00 h on days when browse was not brought to the elephants at the usual time.
Seven additional behaviors were observed and recorded but not included in this report. These included eating, drinking, going into water, sleeping, walking, defecating and urinating. These behaviors were considered biological requirements and are common among all animals. A detailed report on the incidence of these behaviors will be submitted for publication at a future date.
The 14 behaviors described here were also subjected to a statistical factor
TABLE III
Varimax rotated factor matrix for African elephant behavior at the San Diego Wild Animal Park
Variables Factors Commun- _- ~~ ality 1 2 3 4
(1) Trunk to mouth 0.30 -0.42 -0.82 -0.24 0.99 (2) Trunk placement 0.72 -0.07 -0.24 0.20 0.67 (3) Investigative and manipulative 0.86 0.48 0.14 -0.01 0.99 (4) Reach across moat 0.62 0.67 -0.24 0.30 0.98 (5) Trunk intertwining 0.14 -0.32 0.91 -0.17 0.98 (6) Lying down 0.83 0.35 -0.02 0.41 0.98 (7) Eating objects -0.01 - 0.90 -0.26 0.03 0.87 (8) Knocking trunks 0.86 -0.22 0.18 0.33 0.93 (9) Temporal gland secretion 0.51 0.65 0.21 0.45 0.93
(10) Dusting 0.49 0.42 0.65 0.31 0.94 ( 11) Mudding 0.28 0.24 0.05 0.92 0.98 ( 12) Agonistic 0.07 0.93 -0.05 0.31 0.97 (13) Rubbing 0.81 0.51 0.25 -0.08 0.98 (14) Vocalizations 0.30 0.82 -0.13 0.38 0.93
-_
Percent of variation 51.4 17.7 15.4 9.4 - ~_______
268
analysis. The varimax rotated factor matrix for elephant behavior is shown in Table III. Factor 1 indicated high loadings with such activities as trunk placement, investigative-manipulative behavior, reaching across the moat, lying down, knocking trunks, and rubbing. These behaviors accounted for 51.4% of the variation. Factor 2 indicated high loadings with such behaviors as reaching across the moat, eating objects other than food, agonistic be- havior, and vocalizations. These accounted for 17.7% of the variability. Factor 3, indicated high loading with such behaviors as trunk to mouth ac- tivity, trunk intertwining, and dusting. These behaviors accounted for 15.4% of the variation. Factor 4 showed a loading with only a single vari- able - mudding. This behavior accounted for 9.4% of the variation. The four factors accounted for 93.9% of the variation.
DISCUSSION
Female African elephants in captivity demonstrated a variety of interest- ing activities. Their repertoire consisted of 21 different behaviors. These are discussed in terms of categories such as: (1) social behaviors; (2) individ- ual behaviors; (3) biological behaviors; (4) dominance hierarchy; (5) factor analysis.
(1) Social behaviors
Six behaviors were performed with at least two elephants in relationship; these included trunk-to-mouth, trunk intertwining, trunk placement, trunk knocking, vocalizations, and agonistic behavior. This would support the notion that elephants are social animals.
Trunk-to-mouth behavior, which was the most frequently- occurring be- havior among the elephants, played an important role in their learning ex- periences and development of their social life. This activity gave one ele- phant information as to what another elephant was eating or drinking. Douglas-Hamilton and Douglas-Hamilton (1975) describe how baby ele- phants learned what vegetation to eat by sampling what the mother was chewing on. Trunk-to-mouth behavior also appeared to be a gesture of recog- nition and friendship. The elephants engaged in this behavior when they passed each other as they walked about the enclosure, and when an elephant joined another elephant or group of elephants. Trunk-to-mouth behavior also appeared to give assurance and solace to a disturbed elephant.
Trunk intertwining was a preferential behavior. Only one calf had an en- counter with one of the young adult elephants. This behavior was primarily restricted to peer groups. This interaction is interpreted as a form of play behavior and it occurred more frequently among the calves than the young adult elephants.
The elephants engaged in considerable fondling and caressing each other in the form of trunk placement behavior. Contact with the trunk was
269
made to various parts of the body of another elephant. Wankie, the youngest and smallest of the calves, received trunk placements from all the other elephants and more times than any other elephant.
Certain pairs of elephants engaged in trunk knocking behavior more often than others. It is interpreted as a form of play-fighting in which elephants display their exuberance and test their strength against each other.
Six different kinds of uocalizations represented a mode of communication among the elephants. Growls, which were the most frequently occurring sounds, appeared to be a form of recognition or greeting when the elephants approached or passed each other. They were also emitted as a form of aware- ness of environmental activity, such as when the elephants were called by name by the trainer, when trucks were driven near the enclosure or when workmen appeared in the vicinity of the elephant enclosure. Trumpeting appeared to have at least three different meanings. First, this sound accom- panied aggressive behavior such as striking the steel gate with the trunk or body when the elephants were out of food, or when the usual admission time to the barn in the evening was delayed. This sound was also emitted during charges at no apparent object, and it was accompanied by out-spread ears and circular movements. Under these conditions, trumpeting was apparently intended to frighten an adversary. Secondly, trumpeting appeared to be a call for help. On several occasions when this sound was emitted by one of the calves the other elephants rapidly approached that elephant and they would form a defensive circle around her. Thirdly, trumpeting was a sound apparently acknowledging a trumpet sound from another elephant. The African elephants frequently trumpeted in response to trumpeting of Asian elephants in a nearby enclosure. It would be interesting to subject these seemingly similar sounds of trumpeting to a sound-spectrograph analysis. Such an analysis would probably show an actual difference between the trumpetings under different conditions when the sound was emitted. Crying-out appeared to be a sound of fear or pain that was emitted when a calf elephant was being physically attacked by one or both of ,the young adult elephants. The significance of roars, rumblings and squeaks were not ascertained because they were emitted infrequently. These vocalizations are an important means of communication among the elephants and they may be correlated with the head and trunk positions described by Kuhme (1962).
Sounds reported to have been heard by African elephant-hunters are the so-called “tummy-rumblings” (Sikes, 1971). This sound is not respiratory in origin and therefore not included among the vocalizations. It is claimed that this sound is produced when elephants are feeding peacefully and thus it is supposed to be an indication of undisturbed contentment. It is supposed to cease at the slightest intrusion. This sound was not heard among the ele- phants in this study.
Agonistic behavior was observed in which either one, or both, young adults attacked one of the calf elephants. No physical injury was sustained by any of the participants. At the time this agonistic incidence was observed,
270
it appeared to be unprovoked. However, it was later learned that the calf elephant had been removed from the enclosure some time earlier and sub- jected to a variety of training procedures in which the trainer made bodily contact with the elephant. It is speculated that perhaps the human odor on the calf elephant provoked the attacks on her when she was returned to the enclosure.
The six behaviors discussed above indicated that female African elephants are social animals. They apparently had a need for bodily contact and physi- cal interaction with other members of their species. Contact with older ele- phants provided the calves with opportunities to learn patterns of elephant behavior and to test their own individual capabilities through interaction in the form of play, mutual caressing, and non-injurious play-fighting. These findings suggest that in keeping elephants in captivity, housing facili- ties should permit physical interaction among these animals.
(2) Individual behaviors
Eight behaviors have been classified under this heading. Dusting, mudding and rubbing are interpreted as skin-care behaviors. Bere (1966) stated that although elephants are referred to as pachederms, meaning thick skinned, their skin is surprisingly sensitive to touch and temperature variation, and that the skin is well supplied with nerves and blood vessels close to the sur- face. Sikes (1971) has observed dusting and mudding in wild elephants and she believes that the dust and mud-packs insulate the skin from the sun and reduces the irritation from flies and other insect pests. Our observations support such an interpretation.
The elephants frequently resorted to dusting when acacia browse was not brought into their enclosure at the usual time and there was nothing else available to eat. Dusting was also engaged in when the elephants were de- tained from entering the barn at their usual time in the evening. In the barn they were fed hay and other supplements after being chained-up for the night. In these cases, dusting is interpreted as a form of displacement be- havior. Dusting also had emotional connotations. It was significantly corre- lated with temporal gland secretion which we attribute to stress and excite- ment.
The mud and dust formed a thick dry crust over the skin of the elephants. After a period of time this skin-covering apparently irritated the skin or perhaps caused it to itch, and the elephants sought physical objects on which they could rub or scratch various parts of the body, especially those parts that were inaccessible to the trunk. Elephant enclosures should contain low- level objects such as tree stumps or concrete structures that would provide an opportunity to rub the underside or back of the body to relieve itching and irritations of the skin. Our observations of wild elephants in their natural habitats have revealed that they utilized tree stumps and termite mounds as structures to rub against.
271
Rubbing behavior also revealed that elephants were capable of using tools. They have been observed rubbing under their ears and other parts of the body with hay or small pieces of wood held in the tip of the trunk.
With reference to the subject of skin care, we never observed the elephants voluntarily entering the large body of water in the enclosure. Its capacity was such that it could easily accommodate all the elephants at one time. The elephants were persuaded to enter the pond by the caretakers and trainers at least 18 times during the period of observations. We suggest that their reluctance to enter the water voluntarily, even on warm days, was be- cause the same water was used for drinking. Apparently, these elephants preferred not to bathe in the same water they drank. We have noted in the wild that elephants did not enter the water troughs where mechanical pumps raised the water from wells, although the troughs were large enough to accommodate several elephants at one time. The elephants did enter the body of water that accumulated from the over-flow from the trough.
Since dusting, mudding and rubbing are natural behaviors of elephants, facilities in captivity should provide the opportunity for the animals to en- gage in these behaviors.
An unusual behavior was reaching across the moat to obtain flowers and other vegetation and eating it. To reach across the moat, the larger elephants placed their forelegs on the sloping side of the concrete moat and their hind legs on the ground above the moat. This posture placed the elephants in an unusual and precarious position. Reaching across the moat occurred mostly at one place of the enclosure where the moat was least wide and vegetation on the distal side readily accessible.
When the older and larger elephants were engaged in reaching across the moat, Wankie, the youngest and smallest calf, was unable to accomplish this behavior successfully. However, she remembered a pathway into the moat which she discovered during her wanderings about the enclosure. This entrance was located approximat& 40 m from where the other elephants were reaching across the moat. Wankie would immediately walk briskly away from the elephants, enter the moat and walk inside it back to the location where the other elephants were reaching across the moat. From inside the moat she could reach over the moat wall further and more successfully than the others who were reaching across it. The point to be made here is that by taking an extended and roundabout path away from the goal in order to reach the goal successfully is a good example of elephant intelligence and their memory for adaptive behavior. Reaching across the moat also demon- strated the remarkable agility, balance and ingenuity of the elephants.
The elephants were extremely inquisitive animals, as indicated by their investigative and manipulative behavior. The elephants used their trunk not only to sniff odors but physically to investigate and manipulate any objects they could touch with the trunk. They placed pieces of wood, small tree branches, leaves and rocks on to their head and/or back repeatedly as the objects fell off. Anything new in and around the enclosure was thorough-
272
ly investigated by smell and the touch of the trunk. The high incidence of this behavior reflected the elephants’ orientation to their environment, and a need for stimulus-change, and exploration, and especially for objects that could be manipulated. We recommend placing several objects in the enclosure so that elephants can investigate and manipulate them; preferably moveable objects, such as wooden logs, large concrete balls, or truck tires.
Eating objects other than customary food was a surprising and unexpected finding. The calves, more often than the young adult elephants, participated in this behavior.
The explanation for eating objects other than ordinary food may be that since the calves were taken from their mothers in the wild at a very early age, they may not have learned proper feeding habits. Alternatively, eating sand, soil and stones may be an instinctive habit that provides elephants with certain essential minerals which they may not be able to obtain from their regular diet. The cause of coprophagia is uncertain, but we offer some tentative explanations. Since the elephant digestive process is only 35-40’S efficient, considerable undigested matter appears in their droppings. Eating excrement may be palatable and rewarding or at least filling. Another ex- planation of coprophagia may have been the need for establishing a balance of intestinal bacteria. It is also not improbable that the elephants were underfed, and this may have been one form of behavior indicative of that condition. The incidence of coprophagia among wild elephants is not known.
The enormous size and weight of the elephants did not inhibit them from lying down. The elephants did not all lie down at the same time. They lay down to sleep and to cool the body. When one or more specific elephants didlie down, another elephant would stand near the recumbent bodies, usually with the posterior portion of its own body touching an elephant on the ground and the head facing away from the elephants on the ground (Fig. 4). Apparently this is a protective posture by the standing elephant. Carring- ton (1958) described a similar formation of a “protective circle” around the very young when a herd of wild elephants were disturbed or frightened by known or unknown causes. Here the posterior portions of the adult ele- phants were placed together and their heads were facing outwards. We have also observed during our research on wild elephants, that when an elephant dropped to the ground after being hit with an anaesthetic dart, one or more of the unaffected elephants would invariably stand by the one lying down and these had to be chased away before the recumbent one could be approached.
The temporal gland secretion of elephants has been an enigma to scientists. There is considerable disagreement as to the function of the temporal glands in both the African and Asian species of elephant. Most of the explanations offered by Pocock (1916), Bere (1966), Laws (1970), Sikes (1971) or Buss et al. (1976) are speculative. Our behavioral data indicated that the temporal glands secreted under conditions of stress, fear and/or excitement. Whether elephants obtained any information as to the emotional state of another
273
Fig. 4. When a particular elephant lay down, another elephant would stand in the position shown above, alongside the recumbent one.
elephant from the secreting temporal gland was not clear from our study. We did not observe any significant change in the behavior of the other ele- phants when the temporal glands of one or more elephants were secreting. With these behavioral observations, and data obtained in another study in- volving the chemical analysis of the secretion from the temporal glands (Adams et al, 1978), we have tentatively concluded that the secretion from the temporal gland is probably not a mammalian pheromone.
(3) Biological behaviors
There were seven behaviors in this category, which are associated with the usual mammalian biological processes. These include eating, drinking, going into water, sleeping, walking, defecating and urinating. Nothing of unusual significance occurred in the performance of these behaviors. Such parameters as frequency, quantity, duration, etc., would be more appropriately handled in a future report on the physiology of elephants.
(4) Dominance behavior
Sabu, one of the two young adult elephants, was the most active performer of all the seven elephants. She attained the highest frequency in 9 of the 14 behaviors reported. She was second in frequency of two behaviors, and third
274
in one behavior. We therefore declared that Sabu was the dominant female in this group of elephants.
Sikes (1971) had observed elephants in the wild and she concluded that elephant leadership is determined by size, strength and age, and that these characteristics were combined with experience and sagacity. We believe that Sabu has demonstrated these characteristics in captivity.
Based on the amount of participation in the various behaviors described herein, we suggest that the dominance hierarchy of the elephants in this study appears to be in rank order as follows: 1, Sabu; 2, Shapie; 3, Mandavu; 4, Wankie; 5, Madundamella; 6, Hatari; and 7, Deteema.
(5) Factor analysis
Statistical factor analysis of the 14 behaviors revealed four significant factors. The cluster of six variables under Factor 1 are interpreted as curiosity and/or awareness of stimulus change. The cluster of four variables under Factor 2 are interpreted as aggressive activity. The cluster of three variables under Factor 3 are interpreted as pleasurable or playful activity. The single variable with high loading under Factor 4 is interpreted as body comfort. These hypothetical variates describe, summarize and explain the observed behaviors.
In conclusion, this study has revealed that female African elephants have demonstrated a repertoire of interesting social, individual and biological behaviors. We fully recognize that we have not observed the entire repertoire of behavior of African elephants in captivity. The elephants in this study were immature, and all of the same sex. Considering their average life-span to be about 60 years, we obviously have a lot more to learn about the be- havior of elephants in captivity. Similar studies of mature elephants, with interactions of both sexes, may reveal the necessary and sufficient condi- tions under which elephants reproduce in captivity.
SUMMARY
(1)
(2) (3)
(4)
(5)
Seven female African elephants, 2 young adults (ages 14 and 15 years), and five calves (ages 5 years), were observed for a total of 558 hours and 22 minutes during a period of one year. A repertoire of 21 different behaviors was observed. The elephants activities were divided into five categories: 1, social be- haviors; 2, individual behaviors; 3, biological behaviors; 4, dominance hierarchy; 5, factor analysis. The social behaviors consisted of trunk-to-mouth trunk intertwining, trunk placement, trunk knocking, vocalizations and agonistic behavior. The individual behaviors consisted of dusting, mudding, rubbing, reach- ing-across-moat, investigative and manipulative, eating objects, lying down and temporal gland secretion.
275
(6)
(7)
(8)
(9)
(10)
The biological behaviors included eating, drinking, going into water, walking, defecating and urinating. Dominance hierarchy. Based on the frequency of performance of the categories of social and individual behaviors, the dominance hierarchy of the elephants in rank order was Sabu, Shapie, Mandavu, Wankie, Madundamella, Hatari and Deteema. The factor analysis of the 14 activities under categories of social and individual behaviors showed high loading under four factors, namely: 1, curiosity and/or awareness of stimulus change; 2, aggressive activity; 3, pleasurable or playful activity; 4, body comfort. The most frequently occurring behavior was the placement of the trunk of one elephant into or near the mouth of another elephant. Given the opportunity in an environment conducive to variation, African elephants demonstrate a variety of behaviors hitherto unquanti- fied and unreported.
ACKNOWLEDGEMENTS
We wish to thank Dr. Kurt Benirschke and the Research Committee of the San Diego Zoological Society for the privilege of conducting this study. Grateful appreciation is also extended to Dr. James Dolan, Jr., of the San Diego Wild Animal Park, for the cooperation and assistance we received from all staff members during the period of this research.
REFERENCES
Adams, J., 1973. Capture of African elephants. Zoonooz, 46: 8-9. Adams, J. and Berg, J.K., 1974. A preliminary report on the repertoire of behavior of
African elephants in captivity. Paper presented at the Animal Behavior Society Meet- ing, University of Illinois, Chapaign-Urbana, IL.
Adams, J., Garcia, A., III and Foote, C.S., 1978. Some chemical constituents of the secretion from the temporal gland of the African elephant (Loxodonta africana). J. Chem. Ecol., 4: 17-25.
Bere, R., 1966. The African Elephant. Arthur Barker, London, 96 pp. Bolwig, N., 1965. Observations on early behaviors of a young African elephant. Int. Zoo
Yearb., 5: 149-152. Bolwig, N., Hill, D.H. and Philpott, M., 1965. Hand-rearing an African elephant. Int. Zoo
Yearb., 5: 152-154. Buss, I.O., Rasmussen, L.E. and Smuts, G.L., 1976. The role of stress and individual recog-
nition in the function of the African elephant’s temporal gland. Mammalia, 40: 437-451 Carrington, R., 1958. Elephants. Chatto and Windus, London. Douglas-Hamilton, I. and Douglas-Hamilton, O., 1975. Among the Elephants. Viking, New
York. Hediger, H., 1964. Wild Animals in Captivity. Dover, New York, 207 pp. Hediger, H., 1968. The Psychology and Behavior of Animals in Zoos and Circuses. Dover,
New York, 166 pp. Kuhme, W., 1962. Ethology of the African elephant. Int. Zoo Yearb., 6: 113-121.
276
Laws, R.M., 1970. Biology of African elephants. Sci. Prog. Oxford, 58: 251-262. Pocock, R.I., 1916. Scent glands in mammals. Proc. Zool. Sot. London, 2: 742-755. Rensch, B., 1957. The intelligence of elephants. Sci. Am., 196: 44-49. Reuther, R.T., 1969. Growth and diet of young elephants in captivity. Int. Zoo Yearb.,
9: 168-176. Sheldrick, D., 1975. Survival at Tsavo. Africana, 5: 30-35. Sikes, S.K., 1971. The Natural History of the African Elephant. Weidenfeld and Nicolson
London. Young, E. and Oelofse, J., 1969. Management and nutrition of 20 newly captured young
African elephants. Int. Zoo Yearb., 9: 179-184.