Journal of Animal Research & Veterinary Science Category: Agriculture Type: Research Article
Communication in a Small Herd of Semi-Domestic Elephants another Interpretation of their Social Organization, Social Contract and World View
- Marthe Kiley-Worthington1*
- 1 Centre De Eco Etho Recherche, La Combe, Bezaudun Sur Bine, Drome, France
*Corresponding Author:
Marthe Kiley-WorthingtonCentre De Eco Etho Recherche, La Combe, Bezaudun Sur Bine, Drome, France
Tel:+33 475532027,
Email:marthekileyworthington@gmail.com
Received Date: Dec 07, 2018 Accepted Date: Jan 17, 2019 Published Date: Jan 31, 2019
Abstract
Summary
The communication between 6 individually identified free ranging young elephants were observed for over 200 hours (1278 elephant hours) in a nature reserve in Zimbabwe. 97 different behaviours, some explicit (whose meaning is clear) and some implicit (whose meaning is hidden), were recorded. Visual signals were the most common. The meaning of the 22 most common behaviours was assessed from recipient responses. Two surprising results were that (i) many behaviours were ignored by the recipients, and (ii) reciprocity was common. Correlations between the rank orders in the 4 different behavioural categories (aggression, affiliation, avoidance and interest) did not indicate an overall “dominance order”. Rather it indicated that behaviour encouraging group cohesion (showing interest and affiliation) was much more common than any related to competition in the group (that is aggression, avoiding and withdrawing). It is argued that the large number of implicit behaviours which indicate slight arousal also indicate “uncertainty”. Many behaviours ‘meanings were context dependent (i.e. the same behaviour used in a variety of contexts with its particular meaning only assessable from the context). To interpret the meaning of messages in this way, elephants must be aware of others intentions and desires, as well as others’ knowledge and roles in the society. The implication of these results for a different understanding of elephant’s social organization and mental aptitudes are discussed.
2140 interactions were recorded in 213 hours of observation (1278 elephant hours) in a group of 6 individually identifiable free ranging orphan elephants between 10 and 19 years old on a nature reserve in Zimbabwe. 97 defined behaviours were recorded, performed by the initiator or as responses. Visual signals, often associated with other sensory modalities (smell, taste, touch), were the most common. Vocalizations, audible to the human ear accounted for 0.5%, although infra-sound and seismic vibrations were not measured. The explicit and implicit meanings of the 22 most common behaviours were assessed from (i) recipient’s responses, (ii) ethological theory & (iii) folk knowledge. Many behaviours were frequently ignored by the recipient, (14 of the 22 most common behaviours) and the recipient responded reciprocally more often than expected (p<0.05). In addition, ‘interest’ in the performer, and ‘affiliation’ were more common than behaviours associated with conflict or any “dominance order” (p<0.01). Behaviours typically produced in approach/avoidance and frustration situations were common and carried a message that the communicant was “uncertain” what to do. The rank orders in the 4 categories of batched behaviours related to their normative meanings were correlated. This indicated that individual had different roles in the society, but no evidence of any overall” dominance order”. The meanings of many messages were context dependent, conveying information concerning the general level of excitement/arousal with the particular emotional state read from the context. The implication of these findings for further understanding elephants’ social contract, theory of mind, epistemology and evolutionary behaviour are discussed.
Keywords
INTRODUCTION
It has been widely believed that human language has an over powering influence on cognitive [14-17]. As a result, many studies have either focused on assessing whether species can learn to use an artificial human-type, context-independent language [18-22]. Anderson [23], or testing if these species have human mental characteristics such as manipulating and deceiving [6].
Communication in elephants, like that of almost all behaviours of mammals, is flexible and adaptable [24,25]. The specific meaning of the message conveyed by a particular behaviour may be independent of the context in which it occurs, that is “context independent”, similar to much of human language. But it may be “context dependent”, where the meaning conveys the state of arousal of the communicant, but the detailed meaning is read from the context [26,27].
Another feature of elephant’s communication is that it often multi-sensory. A summary of how the different senses are used in communication follows.
Vocal communication has received most attention in a range of species including elephants (e.g. birds, whales, see forthcoming review on audition in mammals). Their auditory sensitivity ranges from infra sound to higher frequencies than humans [28]. Their vocalisations have been studied since the 1960’s [24,25, 27,29,30-32] and a number of studies have focused on the use of ultra-sound by elephants to communicate over several kilometers [33-37] identification of individuals by their vocalizations [33,38,39] and communication by sensitivity to seismic movements has also been demonstrated [40-42].
Olfactory and gustatory communication are often linked but have received less attention in elephants, although they have a very large olfactory sensitive area in the trunk, taste buds on the tongue and around the mouth, and a relatively large olfactory bulb in the brain [1,6,43], mentions that they receive and analyze a great deal of olfactory information which is important to them. The vomeronasal organ attached to the base of the brain is considered to be for the careful investigation of strong, pertinent or unfamiliar smells [44]. It is uniquely large in the elephant and has paired palatal ducts, a folded epithelium and a dorso-medial position of the neuro-epithelium indicating its complexity and links to the cerebral cortex [45], and elephants perform flehmen which is associated with the vomeronasal gland [46]. They have cutaneous glands around the temple and inter-digital glands that produce steroids as well as other secretions and have been found to secrete pheromones related to sexual status. Olfactory research has concentrated on sexual pheromones [45,47-50], but elephants have sweat glands well distributed over the body [51,52], which may produce individual smells. The multi-sensory trunk acts as a hand, a nose and a mouth thus multi-sensory information is inevitably acquired. Taste plays a major role in the choice of diets of course, and frequent tasting another (see below), is likely to be important in communication, but has been little investigated to date.
Some tactile communication has been studied with the salience of the trunk demonstrated [53-55]. Elephants are capable of extraordinarily complex manipulations with the trunk; within the blind area (see below), by touching and presumably smelling, they can sort beans that they wish to eat from those that they do not (personal video), or find rings in piles of hay with their trunks (Circus Knie). Communication by touching different parts of each other’s bodies also are important [53], but such movements also always have visual communicative cue.
Visual Communication has not been very carefully investigated in detail, although explicit trunk movements have been recorded. The first step is to study what they can see. First, they have dichromatic vision, like some colour blind humans [56], but further research is required. Due to the position of the eyes on either side of the large head, elephants have two large simultaneously observed monocular visual fields which enable him/her to acquire visual cues from a large area (300°). But they have a relatively small binocular visual field, approximately 25-60°, compared to the 100-120° of humans. The breadth of the head with the trunk protruding directly in front of the laterally placed eyes ensures that s/he has a blind spot of approximately 2m directly in front of the head [57,58]. Both the placing of the eyes, and the two simultaneous monocular visual field, presents a different visual appreciation of the world from humans’ relatively small monocular, but large binocular visual field.
It has been maintained that elephants have poor vision [51,59,60], although visual signals, such as some trunk movements [24,25,60,61] and head, tail and ear movements [24-26,29,57], have been described. Some visual displays associated with must [62] and female reproduction has also been described although these visual cues may have a biochemical foundation [63].
It is well known that elephant as well as other domestic animals react to slight visual cues indicating intentions that are emotional states. They also show an ability to imitate others [58,64], indicating perhaps emotional contagion [65,66]. These abilities have been used for centuries [67], by their handlers. Thus, such critically assessed information or “folk knowledge” must be part of any good scientific investigations, if we wish to progress in our understanding [68-71], rather than just “discovering” what is already known.
One of the objects of this paper is to report the explicit behaviours in communication (the behaviours whose meanings are clear) and to investigate the responses they cause. The implicit behaviours which have often been ignored in scientific reports, (although their meaning is often known by those who deal with elephants) have also been recorded. First, both the explicit and the implicit behaviours must be defined, their meanings critically evaluated and their involvement with other senses recognized.
Studies of explicit behaviours related to competition between individuals and this often comes with a commitment to outlining a “dominance hierarchies” [72-74]. The inclusion of both afflictive and implicit behaviours in communication might suggest different interpretations of elephant’s social contract.
Non vocal signals are unlike vocalization in their communicative value because vocalizations are only made to communicate [29]. But, visual, olfactory, taste and tactile messages may have other functions and be performed without intent to communicate, even though a message concerning their emotional state may be transferred to others watching. For example when one elephant sees another putting his trunk out to smell a strange object, he receives the message that “there is something interesting there”, even though the behavior was probably performed from curiosity, rather than with intent to communicate. Thus, the communicative meaning of many messages can be complex.
For many movements and vocalizations, the meaning may act as an analogue system indicating the general level of arousal of the animal so the same behavior may occur in a variety of contexts [26,27,75]. The behavior itself does not convey why s/he is in this state. For example, the elephant shakes his/her head when s/he (i) has an irritation on the head, (ii) is scared, (iii) about to attack, (iv) excited, (v) anticipating something (vi) cannot obtain a desire goal. In essence, the animal indicates that it is aroused, but “uncertain” what to do next. This has been demonstrated for many vocalisations, including the elephant rumble [27], head and tail movements in ungulates, elephants and carnivores [26,29,30]. The cause of the arousal can be traced to frustration or approach avoidance conflict [26,30,54,76-80].
Whether the individual is interested, aggressive, and friendly or is experiencing other emotions such as curiosity, fear or embarrassment [81], will be read from the context. Thus, a better understanding of visual signals and the consequent mental aptitudes could further illuminate the elephants’ social contract.
Undeniably, such a study is difficult. It requires experienced observers who attend to details, are inter-observer reliable, familiar with the species and can recognize individuals. Different combinations of behaviours may be performed simultaneously so some behaviour may be missed. Although not all the olfactory messages, seismic vibrations or infra-sounds can be perceived by the human observer without special equipment, if information has been transferred it is likely that there will be some visual cues.
To date, there has been no quantitative analysis of the meaning of many of the visual messages conveyed by elephants, and the importance of their trunk for manipulation and multi-sensory communication needs further study although see Kiley-Worthington and Plotnick et al., [61,82]. As Langbauer et al., mention “more work needs to be done to elucidate the specific role of many signals”.
AIMS OF THE STUDY
(1) To assess the importance and multi-sensory role of visual signals in elephant communication
(2) To ascertain which behaviours are most common, both from the initiators (performers) and as responses.
(3) To help derive the meaning of different behaviours from the analysis of responses
(4) To suggest meanings for implicit visual messages
(5) To correlate the rank orders between the different behaviours and outline the implications of these results founder standing elephant social ontology, their social contract and mental aptitudes
(6) To consider the evolutionary importance of this.
Elephant communication shown in figure 1.

METHODS
Subjects
Husbandry and daily routine
Observations, training of observers & inter-observer reliability
Behaviour |
Definition |
1. Ignore |
Make no response to behaviour directed at the recipient for 20 secs afterwards. |
2. Approach /make contact |
Walk directed towards another for >5steps to within 5m |
3. Contact activity |
Stay within 1m for at least10sec while walking/running |
4. Contact break |
Walk away from another for at least 10m |
5. Turn body towards |
Whole body and head turned towards another and take at least 3 steps. |
6. Follow |
Follow an individual for at least 10m |
7. Turn head towards another |
Turn head only towards another |
8. Trunk extend towards another |
Trunk held out towards another for at least 5 secs. |
9. Tail wag |
Slower sideways movement of tail when no obvious flies were around |
10. Tail swish |
Rapid move of the tail sideways at least 5 times |
11. Tail up |
Tail raised up above the horizontal and held for at least 5sec |
12. Tail withdraw |
Tail pulled tight over anus |
13. Flap both ears forwards & back |
Move ears back and forth at least 3 times. |
14. Both ears back |
Ears compressed against the head with visible muscle contraction |
15. Ears pricked |
Both ears forward, attention directed at something held for at least 10sec |
16. Swing trunk |
The hanging trunk is swung back & forth or right & left at least 5 times. |
17. Trunk up |
Trunk raised above head |
18. Trunk down & relaxed |
Trunk hanging usually on ground |
19. Trunk forward investigating object |
Trunk held forward to smell the object of attention better |
20. Puff |
Air released from trunk with a puffy noise, less abrupt & loud than “blow” |
21. Blow |
Sudden blast of air from trunk giving rise to sudden abrupt noise. |
22. Rumble |
Low amplitude and pitch rumbling vocalisation |
23. Trumpet |
Loud higher amplitude vocalisation with changing pitch |
24. Head shake |
Lateral shaking of the head at least 3 times. |
25. Head nod |
Up and down movement of the head at least 3 times |
26. Head throw |
Trunk and head thrown up and sideways at least 1 time |
27. Head up |
Head held up high above temple for at least 5 sec |
28. Walk |
Lateral 4 time walk of at least 5 paces |
29. Run |
Diagonal or 4x fast walk for at least 5 paces |
30. Lie on sternum |
Four feet spread back and forwards with weight on sternum |
31. Lie flat |
Lying on side with legs laterally on the ground |
32. Getting up |
Front feet out and heave self up |
33. Wallow |
Lye down and move around in water or mud |
34. Urinate |
Eliminating urine for at least 10sec |
35. Defecate |
Eliminating faeces from the anus |
36. Fart |
Air released through the anus making a noise |
37. Sigh |
Slower than normal large breathe taken in and slowly released |
38. Chew |
Moving lower jaw up and down at least 5 times |
39. Paw |
Moving front leg backwards and forwards touching the ground |
40. Swing leg |
Lifting a front or hind leg and swinging it back and forward in the air at least 3x |
41. Raise leg front |
Holding a leg in the air for at least 5 secs |
42. Raise or rest hind leg |
Resting one leg with weight on the other hind |
43. Stamp front or hind leg |
Lift leg up and down hard and faster than normal |
44. Increase postural tonus |
Head, tail up stand erect |
45. Decrease postural tonus |
Head tail and whole body relaxed |
46. Tusks pushing into ground |
Kneeling and pushing tusks into earth, moving head |
Touch/scratch/ smell self |
|
47. Rub head on object |
Head against object and rubbing up and down at least 3 times |
48. Trunk to own mouth |
|
49. Trunk to own penis |
|
50. Trunk to own foot |
|
51. Scratch or rub self with trunk head & eyes |
Trunk twisted to rub the different area |
52. Trunk on shoulder or ears, |
|
53. Trunk between legs |
|
54. Trunk to feet or legs rubbing/investigating |
|
55. Any other area of body |
|
56. Cross legs |
Stand with front or hind legs crossed |
57. Erection of penis |
|
Scratch/smell/touch other |
Recorded if trunk or parts of their bodies touch |
58. Trunk into others mouth |
Trunk enters open mouth of other and remains in for more than 5sc |
59. Touch neck of other with trunk |
|
60. Touch head of other with body or trunk |
|
61. Touch trunk of other |
|
62. Entwine trunks |
Twist trunks around each other |
63. Nibble other’s body with trunk |
Trunk touching and nibbling with prehensile lips any part of another 5sc< |
64. Touch leg |
|
65. Touch quarters of other |
|
66. Touch other’s penis |
|
67. Touch between legs other |
|
68. Rest chin on another |
Chin rested with trunk often over body |
69. Trunk smelling other |
Not closer than 10cm smelling particular area |
70. Trunk pushing at another’s face or thrown forward |
Either trunk pushing or thrown at face of other |
71. Smell/taste urine of other |
Trunk either in urine or smelling urine on ground |
72. Rub body on another |
Rubbing body against any part of another’s body |
73. Touch tail of other with trunk |
|
74. Smell vulva of other with trunk |
|
75. Smell penis of other with trunk |
|
76. Mount another |
|
Other behaviour to another |
|
77. Tasting others urine |
Trunk to own mouth after being in others urine. |
78. Stand over another |
Standing over another lying down for at least 10 sec |
79. Push another with head or tusks |
Head or tusks pushing another’s quarters or side |
80. Flehmen |
Raising trunk and testing urine probably with vomeronasal gland |
81. Turning back on other |
Turning face away from the other |
82. Grab tail of another |
Trunk holds the tail of another |
83. Circle |
Walk around another in a semi circle |
84. Barge another |
Walk fast or run at and push another |
85. Watch other |
Watching another carefully either with one eye or both for at least 10 secs. |
86. Social Play |
Touching, pushing, sparing, chasing each other without intention to hurt. |
87. Object Play |
Lifting/throwing/ pushing etc object |
88. Push another’s head |
With head or tusks |
89. Tusks to tusks |
Pushing twisting tusks to tusks |
90. Tusks to another part of body |
Tusking any other part of body of other |
91. Tusk to legs of other |
|
Avoid or withdraw from another |
|
92. Turn away |
Turn away from another for at least 5 sec |
93. Turn and walk away |
Turn and walk at least 5m |
94. Back off |
Back away from another at least 5m |
Aggressive acts |
|
95. Chase |
Run towards another head up at least 5 m |
96. Push tusks into another’s body |
Anywhere on the body other than to tusks |
97. Tusk to tusk pushing |
|
Statistical method
Behaviour |
No. Times Exhibited by Performer. P |
No. Times Exhibited by a Recipient as a Response. R |
Total Number of Times Exhibited |
% of Total |
Approach/contact make |
677 |
59 |
736 |
9.9 |
Ignore |
0 |
712 |
712 |
9.6 |
Tail wag |
220 |
309 |
529 |
7.1 |
Contact activity (walk, stand or lie) |
335 |
148 |
483 |
6.5 |
Flap ears |
210 |
216 |
426 |
5.7 |
Contact break (general) |
150 |
256 |
406 |
5.4 |
Swing trunk |
182 |
108 |
290 |
3.9 |
Turn body towards |
111 |
92 |
203 |
2.7 |
Tusk - to- tusk |
103 |
96 |
199 |
2.7 |
Trunk up |
103 |
78 |
181 |
2.4 |
Trunk to mouth |
121 |
54 |
175 |
2.3 |
Follow |
145 |
28 |
173 |
2.3 |
Turn head towards |
75 |
46 |
121 |
1.6 |
Trunk extend |
74 |
47 |
121 |
1.6 |
Head shake |
61 |
51 |
112 |
1.5 |
Tusk to body |
77 |
10 |
87 |
1.2 |
Both ears back |
23 |
62 |
85 |
1.1 |
Entwine trunks |
37 |
41 |
78 |
1.0 |
Head nod |
39 |
36 |
75 |
1.0 |
Ears pricked |
38 |
33 |
71 |
1.0 |
Blow |
30 |
39 |
69 |
1.0 |
Tail up |
27 |
39 |
66 |
1.0 |
For each social interaction the identities of the initiator/performer and the recipient were recorded and all of the behaviours exhibited by both (up to 5 behaviours might be simultaneously performed). The behaviours were recorded either on dicta phones and then transferred directly into the Mini-Tab statistical package for analysis, or a psion event recorder. Additional observations of maintenance behaviour and proximity relations exhibited by the elephants were also noted every 15 minutes using a focal, scan sampling method which are reported elsewhere [64].
The behaviours recorded and their definitions are shown in table 1.
RESULTS
The analysis of the meaning of the messages were restricted to the 22 most frequent behaviours, selected on the total number of times performed regardless if by an initiator or a recipient. These behaviours accounted for 73% of the recorded behaviours. Table 2 lists them and their frequencies.
The use of the different sensory modalities in the interactions
Sensory Modality |
Number of Different Behaviours |
Frequency |
% of Total (7542) Different Behaviours Observed |
Visual only, as far as could be assessed |
36 |
4307 |
58% |
Visual, tactile/gustatory and olfactory |
29 |
1314 |
18% |
Visual & olfactory |
7 |
1553 |
21% |
Auditory - vocal |
3 |
37 |
0.5% |
Other behaviours whose sensory modality was not clear |
27 |
143 |
1% |
Olfactory communication
Gustatory communication
Tactile communication
Auditory communication
There are also a variety of non-vocal noises made by elephants without the larynx. These include blows, sighing, sniffing and puffs, stamping, breaking sticks and so on. Blow and puff accounted for 1% of the behaviours used in social interactions. Environmental noises made by individuals were not recorded.
Visual communication
The use of the trunk
Trunk Movement |
No. Times Exhibited by Performer, P |
No. Times Exhibited by Recipient, R |
Total Number of Times Exhibited, P+R |
% of 7542 Behaviours |
Swing trunk |
182 |
108 |
290 |
3.9 |
Trunk up |
103 |
78 |
181 |
2.4 |
Trunk to another’s mouth |
121 |
54 |
175 |
2.3 |
Trunk extend to another |
74 |
47 |
121 |
1.6 |
Trunk to vulva |
72 |
15 |
87 |
1.2 |
Trunk to penis of other |
56 |
29 |
85 |
1.1 |
Entwine trunks |
37 |
41 |
78 |
1.0 |
Trunk to body of other |
52 |
12 |
64 |
0.9 |
Trunk to face of other |
34 |
29 |
63 |
0.8 |
Trunk to anus of other |
17 |
43 |
60 |
0.8 |
Trunk to head/shoulder of other |
35 |
23 |
58 |
0.7 |
Trunk to tail of other |
36 |
6 |
42 |
0.5 |
Trunk to own mouth |
19 |
19 |
38 |
0.5 |
Trunk to rear of another |
12 |
1 |
13 |
0.2 |
Trunk down |
4 |
4 |
8 |
0.1 |
Trunk to neck of other |
7 |
1 |
8 |
0.1 |
Total |
861 |
510 |
1371 |
18.1% |
Further discussion on the use of the trunk is below.
Analysis of the meanings of different behaviours
Behaviour Performed |
Number of Times Recorded |
Number of Different Types of Responses |
Most Common Response |
Second Most Common Response |
Third Most Common Response |
Approach/contact make |
690 |
76 |
Ignore310 |
Contact break106. |
Tail wag69 |
Tail wag |
359 |
68 |
Ignore73 |
Tail wag 66** |
Contact31 |
Contact activity (walk, stand or lie) |
339 |
56 |
Ignore137 |
Contact 117** |
Tail wag49 |
Flap ears |
335 |
59 |
Ignore73 |
Flap ears 66** |
Tail wag54 |
Contact break (general) |
80 |
39 |
Ignore 60 |
Tail wag11 |
Follow7 |
Swing trunk |
217 |
50 |
Ignore88 |
Tail wag34 |
Swing Trunk 29**. |
Turn body towards |
161 |
52 |
Ignore 40 |
Tail wag27 |
Flap ears18 |
Tusk - to- tusk |
198 |
35 |
Tusk-to-tusk 84** |
Entwine trunks18 |
Ignore10 |
Trunk up |
169 |
57 |
Ignore29 |
Trunk up 14** |
Tail wag11 |
Trunk to own mouth |
141 |
44 |
Ignore41 |
Contact break20 |
Trunk to own mouth 13** |
Trunk extend |
104 |
40 |
Ignore19 |
Flap ears12 |
Tail wag12 |
Entwine trunks |
79 |
21 |
Entwine trunks 28** |
Tusk-to-tusk17 |
Contact break4 |
Tusk to body |
55 |
37 |
Contact break24 |
Ignore14 |
Leap away13 |
Both ears back |
45 |
28 |
Tail wag6 |
Flap ears4 |
Contact break 3 |
Head nod |
56 |
26 |
Ignore 17 |
Tail wag10 |
Head nod 8** |
Ears pricked |
58 |
25 |
Ignore 14 |
Prick ears 8** |
Tail wag8 |
Blow |
74 |
35 |
Tail wag8 |
Flap ears5 |
Trunk to mouth5 |
Tail up |
59 |
26 |
Ignore 28 |
Tail wag 5 |
Ears back 4 |
TOTAL number |
3467 |
826 |
1051 |
554 |
348 |
Approach, the most commonly exhibited behavior (690 times), caused 76 different behavioural responses (Table 5). All the behaviours had a number of different responses (column 3).
The implicit behaviour, tail wagging or swishing, was the second most common response. It was often performed with other behaviours (33.2%). See below for discussion.
Ignore as a response
Performed Behaviour |
Number of Times Ignored |
Total Number of Times this Behaviour is Performed |
Percentage Each Behaviour is Ignored |
Most often ignored (more than 20% of occasions performed) |
|||
Contact break (general) |
310 |
677 |
60% |
Follow |
73 |
220 |
50.3% |
Approach/contact make |
137 |
335 |
45.8% |
Head nod |
73 |
210 |
43.6% |
Contact activity (walk, stand or lie) |
102 |
150 |
40.9% |
Flap ears |
88 |
182 |
37.6% |
Ears pricked |
40 |
111 |
36.8% |
Turn body towards |
10 |
103 |
36% |
Swing trunk |
29 |
103 |
35.7% |
Turn head towards |
23 |
121 |
34.7% |
Tail wag |
73 |
145 |
33.2% |
Head shake |
26 |
75 |
32.8% |
Tail up |
8 |
27 |
29.6% |
Trunk up |
19 |
74 |
28.2% |
Trunk extend |
20 |
61 |
25.7% |
Least commonly ignored (less than 20% of occasions when performed) |
|||
Trunk to own mouth |
14 |
77 |
19% |
Tusk to body of other |
3 |
23 |
18% |
Blow |
4 |
37 |
16.7% |
Both ears back |
17 |
39 |
13% |
The less ignored behaviours are likely to be those with the strongest messages. These include: (i) putting the trunk into his own mouth. (ii) Tusking another’s body, which caused a leap away, but could also be ignored? The least ignored of all were “blow” and “both the ears back”. The most common responses to both were tail wagging, and/or putting the trunk in his own mouth (Table 5).
Why so many behaviours are frequently ignored is discussed below.
Do as done by, imitation or reciprocity (Table 5 italic)
Performed Behaviour |
Total Times Elicited by Performer, Frequency |
Same Behaviour Elicited by Recipient, Frequency |
Same Behaviour Elicited by Recipient, % |
Value of Chi2 |
P Value |
Contact activity |
335 |
117 |
34.9 |
13.96 |
0.001 |
Flap ears |
210 |
66 |
31.4 |
31.08 |
0.001 |
Tail wag |
220 |
66 |
30.0 |
29.87 |
0.001 |
Ears pricked |
38 |
14 |
21.1 |
16.02 |
0.001 |
Head nod |
56 |
8 |
20.5 |
9.03 |
0.001 |
Swing trunk |
182 |
29 |
15.9 |
15.16 |
0.001 |
Head shake |
92 |
8 |
8.7 |
6.61 |
0.025 |
Trunk up |
169 |
14 |
8.3 |
8.46 |
0.005 |
Tail up |
27 |
3 |
11.1 |
22.45 |
0.001 |
Trunk to mouth |
121 |
13 |
10.7 |
10.33 |
0.001 |
Both ears back |
23 |
2 |
8.7 |
11.08 |
0.005 |
Blow |
30 |
2 |
6.7 |
1.69 |
NS |
Turn head towards |
75 |
3 |
4.0 |
5.01 |
0.025 |
Contact break (general) |
150 |
5 |
3.3 |
5.82 |
0.025 |
Tusk to body |
77 |
2 |
2.6 |
1.09 |
NS |
Approach/contact make |
677 |
17 |
2.5 |
9.92 |
0.001 |
|
|
|
|
|
|
Why reciprocity is so common is discussed below.
Batching the behaviours according to their meaning
Much implicit behaviour are originally related to skin irritation, such as tail wagging, head nodding or shaking, scratching, and ear flicking (further discussion below). These behaviours were frequent, but their meaning in social interactions is not obvious. These results indicate that these behaviours show some arousal because of indecision/ uncertainty of what to do (approach or avoid: frustrated). Therefore, they are placed in the 4th category labeled “uncertain” or indecision. Category 5 is behaviours that occurred too in frequently to analyze.
To summarize, the batched categories are:-
Socially positive behaviours
Socially negative behaviours
Uncertainty or indecision
Unclassified behaviours
The behaviors in each category are listed in table 8. Table 9 shows the frequency that the different categories were recorded in the interactions.
Interest |
Affiliation |
Aggression and Avoidance |
Uncertainty |
Unclassified |
Approach |
Rest head/neck on other |
Barge into other |
Ears Flap |
Rest leg |
Batched Behaviour |
Total Performed |
% Total Performed |
Total Recipient Responses RR |
% of Recipient Responses Recorded |
Total |
|
Affiliative |
1569 |
30.51** |
1018 |
21.47 |
2587 |
24.27 |
Interest |
1341 |
26.77 |
1104 |
21.24 |
2445 |
22.94 |
Uncertainty |
1301 |
25.3 |
1818** |
34.98 |
3119 |
29.27** |
Aggression |
808 |
15.1 |
368 -* |
0.7 |
1176 |
11.2 |
Avoid |
123* |
2.3 |
648 |
1.24 |
771 |
7.2 |
Unclassified |
403 |
7.26 |
136 |
0.26 |
539 |
5.03 |
TOTAL |
5545 |
100 |
5092 |
100 |
10,637 |
99.91 |
DISCUSSION
Comparative Sociality in mammals measured from the frequency of their interactions
Protective responses
The use of the different sensory modalities in communication
Olfactory communication
Information could be acquired in this way about both the past and the future from smells of seasonal changes. For example, Namibian elephants trek around 150km across the desert to baobab trees when they are fruiting, so they must know when this is happening perhaps partly by recognizing smells of seasonal changes. In social contexts, the age and sex of an individual can be recognized from urine or faces and elephants must be able to recognize individual elephants as well as other species by their smells. Indeed, elephants can been taught to discriminate between the smells of familiar and unfamiliar humans, and to track the smell of an individual.
To summarize, smell information will include where another individual has been, what s/he has been eating, who they are, what their sexual status and age is, who they have been with and so on. In effect, for elephants, the olfactory world could be “something like” reading a newspaper for humans, acquiring information about the past (on the wind, in the water, on the earth, from faeces and urine, from tracks and rubs on vegetation), and possible futures (e.g. oncoming oestrus or a possible birth, movements of groups and forth coming musth). Elephants may also be acquiring information concerning other individual elephants’ (and other animals’) moods (e.g. whether frightened, anxious, or relaxed from body odours such as sweat). Particularly interesting smells are also indicated visually by movements of the trunk.
Gustatory communication
Tactile communication
Auditory communication, audible to the human ear
Visual communication
The trunk, a multi-sensory communicative organ
The trunk is also a manipulative organ, like humans, elephants are manipulators, humans with their hands, elephants with their trunks. All of their food is collected using the trunk, which sorts, selects, prepares and processes it [24,25,58] .The trunk has approximately 500 muscles controlling its movement [57]. Consequently, it can perform a great variety of movements such as:- Gripping, pushing, twisting, pulling, grabbing, touching. It can be turned and twisted into small places, used to investigate, to caress, push or slap other elephants and humans [59]. It is particularly used to investigate others’ body areas rich in smells, such as the mouth, ears, feet, temporal glands, anus and vulva. It can be used with great skill to lift, pick up, balance, carry, place and stack heavy objects and the tusks, head and trunk working together can act as a fore-loader. Adams [67], personal observations in India with timber elephants). It can also be used to find and lift objects under water (personal observations) and find small hidden objects (such as rings and buckles hidden in hay) and is skilled at sorting objects, even out of sight. It is used for breaking or pushing things over, such as trees or fences (including discriminating between electric and insulated lines, in order to lift posts our of the ground [89]. It is used to make and use tools [89,90] and can be used to manipulate objects in a mirror [91], which humans find difficult. The trunk is used as a resonating cavity for vocalisations, and used to learn to imitate human words [92] and invent new sounds (personal videos elephants in the [93].
The use of the trunk in communication accounted for 18% of all the behaviours in interactions. The 16 different trunk movements had meanings inall 4 of the batched behavioral categories. The trunk is also used of course, for self directed behaviours such as self grooming, smelling, rubbing, scratching, stroking, and essential for drinking, food selection and processing.
The trunk placed in the performer’s own mouth was frequent in training experiments [94], occurring when the individual was uncertain what to do, or confused. It occurs in the same situations in which children put their finger or thumb in their mouths which demonstrate uncertainty [95]. On several occasions when teaching elephants with unprotected contact, when they were frustrated or annoyed, they stamped one front leg. This is a similar behavior given in a similar situation by a child stamping a foot. Beating the trunk on the ground is another well known behaviour in captive elephants associated with frustration (e.g. awaiting food or to be unshackle [51], but was not observed during these observations.
One of the most interesting movements is the trunk placed in another’s mouth, a relatively common behaviour (performed 121x’s by an initiator and by a recipient 54x’s: Total 175x’s). Here the performer acquires information about the recipient and it also demonstrates “trust/confidence/amicability” as biting the other’s trunk would be easy. This action has frequently been observed to have a calming influence on the recipient (observations & videos). Some handlers imitate this action by stroking the tongue of an elephant when greeting. The establishment of predictability in behaviour, “trust” or “confidence in each other” has been widely used by good animal handlers for generations [85,96-98] and is now considered to be one of the most important components in establishing good cooperative animal handling and teaching [58].
The hand/nose/tongue trunk is used in almost every type of situation to manipulate the world, to communicate with others and to read intentions and emotional states of others. Without being aware that his trunk is HIS, the elephant would be unable to do these things, indicating his self awareness of his trunk and his ability to control what, where and when it moves. (Further discussion below).
The indication of parallel experiences between the elephant trunk and human hand underlines the importance of taking a conditional anthropomorphic approach to further comprehend elephant epistemology and mental aptitudes [68,69,71,82].
The meanings of the different behaviours
The meanings of many behaviours are self evident; that is, they are “explicit” and can be batched into two categories (i) aggression and avoidance / withdrawing all of which will tend to split the group. (ii) Affiliation and interest in the other, which will encourage group cohesive or sticking. The third category contains behaviours frequently used in communication, which have an “implicit” (inherent, hidden) meaning. Many visual signals are in this category. Further discussion follows.
“Uncertainty”. Behaviours indicative of approach avoidance conflict and frustration, convey a message of indecision
An interesting behaviour in the same category is placing the trunk in his own mouth. Children may also place their fingers in their moths when in indecisive situations [26,29,58,95], another similarity to humans.
Anselme & Gurturkun [104], in a recent paper shows that “uncertainty” magnifies food searching behavior in birds which then increases exploration and survival. This “uncertainty principle” could equally be applied in communication to reduce potentially emotionally inflammatory situations, encourage group cohesion which in turn will encourage social exploration and curiosity, leading to the acquisition of more environmental and social knowledge.
Ignoring directed behaviours
Therefore, the most likely explanation is that the recipient chose to ignore that behaviour directed at him/her. Detailed investigation of what behaviours were ignored indicates that some of the most frequently ignored demonstrated “uncertainty” (e.g. tail wag, head nod, flap ears, swing trunk and head shake). The message of “uncertainty” or indecision had been received, but, because there was no threat to social cohesion, there was no need for a response. Ignoring in this situation may help to deflate any emotional conflict.
It is less clear why social behaviours, such as approach and contact make, turn body towards, turn head towards, tail up (indicating a rise in excitement), trunk up, and tusking another’s body are so frequently ignored. But this may indicate that ignoring a situation in which conflicts might otherwise arise and grow, will aid group cohesion.
Behaviours that are less ignored will have strong messages. These include putting the trunk in the mouth of another, which shows interest, familiarity, confidence or “trust”. “Blow” and trunk extended sniffing were also rarely ignored both indicating something like” take heed, there is something interesting around”. Drawing both ears back against the head was also infrequently ignored; which may have a message something like “I am slightly concerned about the situation” and the recipient reciprocating message will mean: “I have seen you and am therefore also slightly concern” .
Do as you have been done by/ reciprocity or imitation
Reciprocity and its role in fostering bonding and cooperation has been discussed in some animals [108], but has not previously been reported in elephants. Sixteen of the most commonly recorded behaviours all of which demonstrated affection/sticking behaviour or uncertainty, were very significantly imitated by the recipient. These were those involving orientation movements and postures (ears prick or turn head towards the other,) investigation (trunk up & smelling), interest in another (approach and contact make). They included those indicating an increase in arousal and preparation for action such “tail up” [26]. Behaviours indicating “uncertainty”/ indecision, were also significantly reciprocated (such as tail wag, head nod, swing trunk, head shake and trunk to own mouth).
It may be that reciprocal behaviour is more common than has been previously reported in many species. Randle & Kiley-Worthington showed that inter-species imitation (the animal imitating a novel action of the human teacher) is frequent in teaching situations and, as in children, can be used as a teaching aid. The function may be to indicate to another; something like “I am also feeling the same way”, that is emotional contagion [65,66], which is another way of indicating awareness of others’ feelings and intentions. It is interesting that elephants can also be taught to imitate humans speech which is suggested helps them to bond with humans [92].
Kingsley [105], maintains that both “do as you would be done by “that is be nice to each other, and “be done by as you did” (if nasty, it will be returned) are moral rules which children should obey. Following on from this, reciprocating with aggressive behaviours when aggressed would ensure that the social contract is adhered to and those who do not adher can leave the society, so the society can continue. Reciprocity of behaviour (and its accompanying emotions) that might threaten the group stability and the social contract: “Be done by as you did”, would then discourage further contravention of the social contract.
Context dependent and context independent meanings
One of the particularly interesting consequences of context dependent messaging is that, because a particular behaviour acts as an analogue system to convey a message about the general level of excitement or arousal of the individual, many behaviours are used in a variety of situations. The specific meaning, that is why the elephant is aroused, can only be gathered from the context, thus whether s/he is frightened, worried, feeling slightly aggressive or sexy will only be gathered from the context. As a result, each individual must be acutely aware of the environment and the intent of others or he will not be able to interpret accurately the meaning of the message. Any witness will have to have an understanding of the situation and be aware of others’ intentions that are his emotions, desires and needs to interpret the message. Such a reading of intentions of others has been labeled having a “theory of mind”: Defined as “attributing perceptual, epistemic, and volitional states to others and themselves” [109-117]. Therefore, animals that use context dependent communication must have a ‘theory of mind’ [71,110] or they would not be able to communicate, have a social contract and a social organization.
By contrast, to understand the meaning of context independent messages (such as human language), it is not always necessary to have an awareness of others intentions, desires and needs because the meaning of the message is constant, whatever the situation, emotions and intentions of the communicant which is what allows human language to be understood out of the present context: On the computer or telephone. This is because context independent communication conveys complex, symbolic non-emotional messages entirely divorced from the context and, consequently, can be understood without having any theory of mind of the communicator and his intentions.
Context dependent communication therefore presents a different complex cognitive assessment of the world from context independent language. But, judgments concerning mental development and complexity, are usually made only by comparison to humans, often because of the overruling importance of context independent language in which humans both speak and think most of the time which acts as a cognitive handicap when attempting to understanding the ontology and epistemology of species who use context dependent communication and who, to understand the message clearly, must be acutely aware of the social and physical environment around them and the feelings and intentions of others [66,71]. Without an awareness of others feelings and intentions, such communication will not function and without communications, there could be no social contract, recognition of other individuality, and their knowledge or roles, that is no “social networking” [11].
Rethinking elephants social contract and their epistemology related to these results
In addition in species such as large herbivours where resources are usually widespread and available for all or none, there is little need for constant competition within the group. Individuals have different personalities and consequently different placings in the different behavioural hierarchies, that is some are more aggressive than others, some more affiliative. Some more submissive, and some more socially involved [85]. But, there is not one individual who is top of the aggressive hierarchy and low in the avoidance hierarchy as there would be if there was a “dominance hierarchy” controlling access to resources. The organization of this society is more complex with individuals having a variety of different roles.
In large free living herbivors since there is no need for constant competition for resources, behaviours and their accompanying emotions which encourage group cohesion would be likely to be common and selected for such as those demonstrating interest and affiliation to keep the group together. Demonstrations of indecision that is “uncertainty”, “reciprocity” and ignoring directed behaviour may also help to here the group by reducing negative emotions and possible group splitting.
The question remains why group cohesion is so important. To grow, survive and reproduce, each individual must become a natural botanist, zoologist, ethologist, meteologist, geologist and geographer that is s/he must become a good “natural ecologist” [81,123]. There is an enormous amount of information to acquire by each infant in order to survive, so perhaps, the most important reason for group living is to enable the acquisition of such necessary knowledge by facilitating observational, social learning and imitation, which aid in a rapid and less risky accumulation of knowledge than learning everything by trial and error. However, it is necessary to know who to learn from: To know what knowledge other individual have. Thus an awareness of the age, personality, role [124] and knowledge of each group member will be crucial. Thus, older animal’s greater knowledge is recognized so they often become decision makers or “matriarchs” [25,71]. Strong, askilled manipulators will have a role of helping infants out of mud holes [125,126].
If the society is to prosper and reproduce, the society’s rules, that is, the social contract must be obeyed. Such rules as “do not attack infants”, ”recognize others roles” or “do not perform socially negative behavior unless another breaks the rules”. If an individual does break the rules, then action must be taken. This would account for the frequency of reciprocal behaviours: “If you are nice to me, I will be nice to you; if you are nasty, I will return it in kind”.
There are “other ways of seeing” outlined by Goethe [127], which may be imperative when considering other species epistemology. One idea that is becoming increasingly apparent and may be one of the most significant and important developments for life, is the recognition of how wide spread symbiosis is, both between and within species [128-130]. The importance of symbiosis in the social world of different species [131,132] and different strategies to encourage this in the social contract is another way forward to an understanding of that species epistemology [66,71, 81,82].
The question remains why, what would appear to be a relatively small evolutionary step, that of associating a particular movement posture or call with a particular message, has not always been taken. But, different cognitive maps are held by different individuals and species, for example blind humans become more sensitive to wind, smells and touch than normal humans, and other “handicaps” in humans also point out different cognitive maps. For different species, an emphasis on olfaction may require a different concept of space and time from that of humans [81]. A different world view will be acquired by two simultaneous large monocular visual fields, compared to a large binocular visual field and poor peripheral monocular vision characteristic of humans. The different world view implicated by context dependent communication also illustrates a different world view since when using context dependent communication, the recipient has to be aware of the “perceptual epistemic, volitional states of the other” to receive the correct message and must “socially network”.
There is much evidence that non-human mammals do recognize other living “beings”; that is they recognize the body/mind/moods/emotions/intentions/knowledge/roles of others, both inter- and intra-species, and discriminate between the living and the dead [25]. Such communication requires an awareness of others body/mind being that context independent communication does not always need. Thus, awareness that the other has a mind is a necessary attribute for context dependent communication.
Many even naïve/wild mammals have an ability to read others feelings and intentions, including those of other mammals, humans. Even naïve rhinos, horses, dogs, cattle, elephants, lions, tigers respond rapidly to humans moods, although some humans may need experience before they are able to read the intentions of an animal [58]. Having an awareness of others and their intentions is a necessary mental attribute for context dependent language users without which they could not have a social ontology, social organizations or a social network. Whether such animals have “reflective self-knowledge” [133-138], is not the same question.
CONCLUSION
Correlations of the individuals rankings in the different batched behaviours, did not confirm that there is any overall “dominant” individual, but that individuals have different roles in the society and that the prime society parameter is to facilitate it’s cohesion: “Sticking behaviour”, not competitive or “splitting behaviour” where some priority of access for an individual that is a “dominance hierarchy” might be the most important organizational parameter.
Positive or “sticking” behavior (affiliative and showing interest in another), are much more frequently than expected which indicates that, for this large herbivore, the cohesion of the group is more important than competition within it (“splitting” behavior).Cohesive and symbiotic behavior, may have been selected for to facilitate knowledge transfer by social and observational learning.
The meaning of much of elephant communication is context dependent that is the same behaviours are performed in a variety of situations. To interpret the specific meaning of a message, the recipient must be aware of the situation and the intention of others. Judgments concerning mental attributes only by comparison to humans discourage consideration of different and complex social ontology’s and can be a cognitive handicap, we should broaden the search in order to better understand a species epistemology. Animals think, and we can begin to understand what different species think about [139,140].
ACKNOWLEDGEMENT
FUNDING
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Citation:Kiley-Worthington M (2019) Communication in a Small Herd of Semi-Domestic Elephants another Interpretation of their Social Organization, Social Contract and World View. J Anim Res Vet Sci 3: 012.
Copyright: © 2019 Marthe Kiley-Worthington, et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
