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Signals, Gestures, and Behavior of African Elephants

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This chapter, which draws on photographs and descriptions in published material to describe many of the displays, signals, postures, and gestures of African elephants, aims to provide a detailed description of the varied elements in the behaviors of elephants so that they can be recognized and contextually understood in the field. Most of the entries in the chapter are true displays or signals. In addition to signals, the chapter includes movements, postures, and gestures that are not necessarily intentful, but which may be a consequence of an elephant's state or situation. Most of these latter behaviors also convey specific information to observers (both human and elephant) and may function as true signals. Finally, the chapter includes a few postures and behaviors that elephants do only in specific situations.
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additional displays and gestures of African and Asian ele-
phants (Poole and Granli [2003], later updated as Poole and
Granli [2009]). Our descriptions were based on the long-
term observations of elephants and inspection of thousands
of photographs and descriptions in published material,
rather than the systematic analysis of video material. This
chapter draws on that body of work to describe many of
the displays, signals, postures, and gestures of African ele-
phants. Its purpose is to provide a detailed description of
the varied elements in the behaviors of elephants so that
they can be recognized and contextually understood in the
eld. In some cases we have supplemented the written de-
scriptions with a photograph, in other cases photographs of
the behaviors may be viewed on the Gestures Database on
www.ElephantVoices.org.
Most of the entries in this chapter are true displays or
signals. In other words, they are behaviors used to trans-
fer information, reducing the observer’s uncertainty about
the actor’s or reactor’s future behavior (Krebs and Davies
1987). In addition to signals, we have included movements,
postures, and gestures that are not necessarily intentful, but
may be a consequence of an elephant’s state or situation.
Most of these latter behaviors also convey specifi c informa-
tion to observers (both human and elephant) and may func-
tion as true signals (e.g., Waiting; Periscope- Sniff). Finally,
we include a few postures and behaviors that elephants just
do in specifi c situations (e.g., Testing- Semen). These may or
may not act as signals to other elephants, but they are inter-
pretable by humans as measures of behavior and are, there-
fore, admissible in a comprehensive ethogram.
Following the convention of Kahl and Armstrong (2000),
A
frican elephants, like other intelligent, long- lived,
and highly social, group- living animals, use a wide
range of sensory channels to communicate with
one another. Visual, tactile, auditory, and olfactory signals
all appear to be important to the social life of an elephant.
Considerable work has focused on both auditory (see
Poole, chapter 9; Berg 1983; Poole et al. 1988; Poole 1982,
1989a; Payne, Langbauer, and Thomas 1986; Langbauer
et al. 1989; Langbauer et al. 1991; Soltis, Leong, and Sav-
age 2005a, 2005b) and olfactory communication (mainly
with captive Asian elephants: Rasmussen [1988]; Rasmus-
sen et al. [1982]; Rasmussen et al. [1993]; Rasmussen, Hall-
Martin, and Hess [1996]; Rasmussen and Schulte [1998];
Rasmussen and Wittemyer [2002]; Rasmussen and Krish-
namurthy [2000]). Surprisingly, few systematic studies have
described the visual and tactile signals of elephants except
where specifi c behaviors explain or reveal underlying pat-
terns of social or sexual behavior (e.g., Douglas- Hamilton
1972; Moss 1983; Poole 1987a).
In the early 1990s Kahl and Armstrong embarked on an
ambitious study to compile an ethogram of the African ele-
phant. In 2000, they named 83 ritualized visual and tactile
displays based on their own observations as well as pre-
viously published descriptions. This initial inventory was
simply a list without description, awaiting the detailed, sys-
tematic analysis of hours of video footage.
In 2002, with input from Kahl and Armstrong and
pending their more detailed study, we developed an online
searchable database, www.ElephantVoices.org, describ-
ing the appearance (written descriptions and photographs)
and contextual use of these 83 displays as well as scores of
Chapter 8 Signals, Gestures, and
BehaviorofAfrican Elephants
Joyce H. Poole and Petter Granli
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110 Joyce H. Poole and Petter Granli
and dust to fl y (Eltringham 1982; Poole 1987a; Payne and
Langbauer 1992; Langbauer 2000). Alternatively, an ele-
phant may frighten a predator (e.g., lions, humans), an ir-
ritant (e.g., egrets, ground- hornbills, warthogs), or an ele-
phant adversary with a Forward- Trunk- Swing, stepping or
lunging forward and swinging or tossing the trunk in its
direction, while simultaneously blowing out through the
trunk (Kühme 1961; Poole 1987a). Similar swinging of the
the fi rst letter of names given for behaviors are capitalized;
if the name contains two words, the fi rst letter of each word
is capitalized and the name is hyphenated. The behaviors
have been divided into several general contexts: Aggres-
sive, Ambivalent, Socially Integrative, Mother- Offspring,
Group Defense, Sexual, Play, and Generally Attentive. Each
of these has been further subdivided. The behaviors are in
bold the fi rst time they are mentioned in the sections where
they are described. Otherwise they are italicized.
Aggressive
Threat
An elephant may threaten another simply by a resolute re-
orienting of its body to gaze at an opponent (Poole, personal
observation; Kahl and Armstrong 2000), or by purposeful,
directed walking toward another. Sometimes just one step
toward another is enough to cause a reaction (Eisenberg,
McKay, and Jainudeen 1971; McKay 1973; Poole 1987a,
1989a, 1999a; Moss 1988, 1992; Ben- Shahar 1999). A
Turn - Toward and an Advance- Toward may be associ-
ated with other aggressive postures such as Ear- Spreading,
Standing- Tall, or Ear- Folding, and so forth.
In an Ear- Spreading threat an elephant faces its oppo-
nent head- on and fully spreads its ears (perpendicular to
its body), thus increasing its apparent size to the viewer
(Kühme 1961; Douglas- Hamilton 1972; McKay 1973). In
a more exaggerated form, an elephant spreads its ears while
Standing- Tall, holds its head held well above its shoulders,
and with tusks lifted, directs its gaze toward its adversary,
as in fi gure 8.1 (Douglas- Hamilton 1972; Poole 1982,
1987a, 1987b, 1987c, 1999a; Kahl and Armstrong 2000;
Langbauer 2000). In an exaggerated form of Standing- Tall,
an individual may stand on a log or an anthill to appear
even taller—a tactic often used by males when they are siz-
ing up one another.
A threatening elephant may press or “fold” the lower
portion of its ears under and back so that a prominent hor-
izontal ridge appears across the ear as depicted in fi gure
8.2 (Moss 1988; Poole 1982, 1987a, 1989a). Ear- Folding
may be used in combination with a variety of other threats
such as Standing- Tall, Advance- Toward, and so forth, to
emphasize that an elephant “means business.” Ear- Folding
that is used in association with Head- Raising, Ear- Lifting,
and / or Rapid- Ear- Flapping is an affi liative, not aggressive,
display and is a typical greeting posture (see Social Integra-
tion, below).
An elephant expresses irritation or impatience with an
individual or circumstance with an abrupt shaking of the
head, or Head- Shaking, which causes the ears to fl ap sharply
Figure 8.1 Standing- Tall, a threatening elephant appears larger by facing its
opponent head- on, with the head well above the shoulders and the ears fully
spread.
Figure 8.2 An elephant threatens another by Ear- Folding, pressing or “fold-
ing” the lower portion of its ears under and back so that a prominent horizontal
ridge appears across the ear.
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Signals, Gestures, and BehaviorofAfrican Elephants 111
This posture may be held at a fast walk or during a Mock-
Charge and / or Real- Charge, especially when the subject
of the charge is of smaller stature. In a sense the aggressor
brings her head and tusks down to her victim’s level (Kahl
and Armstrong 2000). It is one of the more common forms
of high- level aggression between females of different fami-
lies or toward young, non- family males (Poole, personal ob-
servation). Kahl (personal communication) notes that it is
reminiscent of the Forward Threat Display performed by
geese. In a Coalition, two or more individuals work together
to threaten and / or pursue another individual or individuals.
Generally an older female, often the matriarch, comes to
the aid of a younger family member. The helping individual
may approach her companion in parallel; they may vocal-
ize in unison, sometimes clicking their tusks together (Tusk-
Clicking; see below), and then together Advance- Toward
and / or Run- After their adversary with a Bow- Neck display
(Poole, personal observation). In a Group- Charge an entire
family charges toward an adversary en masse in a highly
coordinated manner (Poole 1996; Ben- Shahar 1999). This
behavior is most often observed as an anti- predator display.
Escalated aggression may also involve a variety of
physical contact. An elephant may push its head into the
back, side, head, or trunk of another; such Pushing may
lead to one animal being supplanted, and differs from the
Herding- Push (Lee [1986]; Moss[1988]; Langbauer [2000];
Archie, unpublished data, see below). An individual may hit
or slap another with its trunk, especially older individuals
toward calves. Although Slapping is uncommonly seen in
Loxodonta, it is apparently more common in Elephas (Lee
1986; Kahl and Armstrong 2000). Poking an opponent with
the tip of the tusk, or Tusking, is an escalated form of push-
ing another out of the way. An elephant may tusk another in
competition over resources, or in irritation (Lee 1986; Poole
1987a, 1989a; Kahl and Armstrong 2000; Archie, unpub-
lished data).
A combined highly aggressive form of Pushing and Tusk-
ing results in Ramming, in which an individual lowers its
head with its trunk curled under (to expose tusks) and
rushes toward another, goring or poking it with its tusks,
breaking the skin and potentially causing serious wounds
(Kahl and Armstrong [2000]; Poole, personal observation;
Archie, unpublished data).
Males, particularly those in musth, take physical contact
to an extreme form, rushing at one another and attempting
to gore each other with their tusks, as in fi gure 8.3. Dueling
males tusk and push one another head to head, attempting
to lock tusks and leverage their opponent over or maneuver
him into position where he can be gored. Dueling is associ-
ated with the Head- High, Tail- Raising, Ear- Spreading, Ear-
Folding, Trunk- Out- Stretched (to reduce any blow) (Poole
1982, 1987b, 1987c, 1996, 1989a).
trunk is also observed in play (see Play, following). Lifting
or uprooting objects and throwing them in the direction of
an opponent, or Throw- Debris, is an escalated form of the
Forward- Trunk- Swing. An elephant’s aim can be very ac-
curate even at some distance (Kühme 1961; Poole 1987a,
1987c, 1996; Kahl and Armstrong 2000).
The Head- Nod, a rather jerky up and down move-
ment of the head, which at higher intensities may cause
the trunk to fl op about, is often associated with, or a pre-
lude to, a Mock- Charge (Kühme 1961, 1963; Estes 1991;
Douglas- Hamilton and Douglas- Hamilton 1992; Ben-
Shahar 1999).
Two musth males involved in, or leading up to, an
Escalated- Contest may pace back and forth in a Parallel-
Walk to one another, typically placing a road, log, or other
“barrier” between them (Poole, 1987b, 1987c). Tusking
of the ground and uplifting of vegetation (Tusk- Ground),
or tossing of the head and tusks back and forth through
bushes or other vegetation (Bush- Bashing) creating noise,
commotion, and a demonstration of strength is usually
seen during the maneuvering of two musth males during an
Escalated- Contest (Kühme 1961; Moss 1988; Poole 1987c,
1996; may also be seen in play, below). Escalated- Contests
are usually terminated with the winner tracking the loser
over several kilometers in an aggressive, persistent, and
Prolonged- Pursuit (Poole, personal observation).
Escalation
Lunging and rushing at, chasing after, and physical contact
all characterize more escalated elephant aggression. Ele-
phants may be observed to run after or chase an opponent
in a Pursuit (Poole 1982, 1987a, 1989a; Lee 1986). This
increased aggression on the Advance- Toward takes several
forms. A Mock- Charge is a rushing toward an adversary
or predator, usually exhibiting Head- High (see Sexual Ad-
vertisement below) and Ear- Spreading that stops short of
its target (Douglas- Hamilton 1972; Langbauer 2000); an
elephant may aggressively Forward- Trunk- Swing or Kick-
Dust (see below) as it abruptly stops (Poole, personal ob-
servation). A Mock- Charge is often associated with a shrill
Trumpet- Blast (see Poole, chapter 9). In a Real- Charge
an elephant rushes toward another elephant, predator, or
other adversary with the apparent intention of following
through. The head may be held high or lowered; the ears are
spread and trunk curled under (so that tusks make contact
rst). A Real- Charge is usually silent (Poole 1987c; Kahl
and Armstrong 2002). In the Bow- Neck charge, the aggres-
sor (usually female) lowers her head by bowing her neck
downward and simultaneously tilting her head upward so
that her tusks are approximately horizontal. The Bow- Neck
may be associated with Ear- Flattening and / or Ear- Folding.
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112 Joyce H. Poole and Petter Granli
(Poole, unpublished data). Similarly, around a scarce re-
source or object of interest such as a drinking place, mud-
wallow, fallen tree, or an object of play, lower- ranking indi-
viduals may engage either in displacement activities or stand
quietly as they wait their turn.
When joining a resting group and also in the context of
high- level greetings (see social integration below) an ele-
phant presents its posterior and walks Back- Toward an-
other, usually higher- ranking animal (Moss 1988; Kahl
and Armstrong 2000; Langbauer2000; Archie, unpub-
lished data). A Back- Toward another may be an appease-
ment gesture or sign of respect, especially within families
(Poole, personal observation) or used to avoid triggering
an aggressive response in a larger social context (Kahl and
Armstrong 2000). As an interesting aside, Hyraxes (Pro-
cavia) also Back- Toward in similar circumstances (Croze,
personal communication). Elephants may also be seen to
Back- Toward a dead elephant or to the bones of an ele-
phant (Moss 1992; Poole 1996; and see below).
Retreat
An elephant may avoid confl ict with a threatening individ-
ual by adopting a Head- Low posture, orienting or walk-
ing away, or retreating from, the aggressor while looking
back (Turn- Away, Retreat- From, Look- Back; Poole [1982,
1987a, 1989a]. Such avoidance behavior is typically as-
sociated with a wary Look- Back over the shoulder (Poole
1987a, 1999a; Kahl and Armstrong 2000). An individ-
ual may also Retreat- From an aggressor by backing away
(Back- Away; Kühme [1961]; Poole, personal observation).
During escalated aggression an individual may avoid con-
ict with a Run- Away; this is as a common response to a
Pursuit (Poole 1982).
Post- Confl ict Display
Following an aggressive interaction or more prolonged con-
ict, elephants may engage in a number of post- confl ict be-
haviors. For example, in an apparent sign of bravado an
individual may adopt an Exaggerated- Walk, an obvious
bouncing swagger in which the front legs appear to bend
more, and the movement of the head up and down and the
trunk from side to side is overstated (Buss and Estes 1971;
Hendrichs 1971; Poole 1987a; Moss 1988, 1992; Balfour
and Balfour 1997). A similar walk is seen as any age / sex
comes down a slope, particularly toward water. Croze (per-
sonal communication) refers to this as the “water- walk,”
which he believes may take its form from the inevitable
down- slope toward water. While this may not be a signal,
he has seen it displace subordinate males from the water’s
edge.
The term Escalated- Contest covers all displays that may
be seen in the context of a serious battle between two males.
These may include: Dueling, Tusk- Ground, Parallel- Walk,
Throw- Debris, Bush- Bashing, Trunk- Out- Stretched, Head-
Toss, Trunk- Drag, Trunk- Curl, Run- After, Run- Away, and
others (Poole [1987a, 1987b, 1987c, 1989a]; some de-
scribed below).
Submission
A submissive elephant holds its head low (Head- Low) so
that the top is below the level of the shoulder blades, mak-
ing the individual appear smaller. This posture is often as-
sociated with a Turn- Away and followed by a Retreat- From
(see Retreat below). A Head- Low can be a persistent dis-
play by a low- ranking male just as Head- High can be a per-
sistent display by a high- ranking male (Eisenberg, McKay,
and Jainudeen 1971; Moss 1983, 1988, 1992; Poole 1987b,
1989a, 1996, 1997; Estes 1991; Spinage 1994); the former
is often observed in young males in presence of musth male.
The Head- Low may be an appeasement gesture, which sup-
presses aggression in the other individual. A submissive ele-
phant may also hold its ears fl attened (Ear- Flatten) against
the body (Poole, personal observation; Kahl, personal com-
munication). An elephant normally holds its ears in a re-
laxed position such that they rest approximately 10 to 30
cm from the side of the body. In this relaxed position the
ears do not appear stiff or tense, lifted, spread, or pressed
against the body.
In the presence of high- ranking individuals around
a limited resource (at a fallen tree, or in the company of
an estrous female), lower- ranking individuals (most often
young males) may be observed to make a small arc or
Skirt- Around the group to try to establish a better position
Figure 8.3 Physical aggression takes an extreme form among males in musth.
Dueling—males rush at one another. Rammingmales ram into one another.
Pushing—males push each other head- to- head. Tusking—an attempt to gore
the other with the tusks.
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Signals, Gestures, and BehaviorofAfrican Elephants 113
McKay 1973; Adams and Berg 1980; Douglas- Hamilton
and Douglas- Hamilton 1992; Spinage 1994; Sukumar
1994; Kahl and Armstrong 2000). Alternatively, an indi-
vidual may begin to pluck at vegetation, as if foraging, but
may not actually ingest any of the material. Even if the ele-
phant does eat, it does so in a halfhearted or distracted fash-
ion and may slap the vegetation against foot or other part
of own body and then discard it (Kühme 1963; Eisenberg,
McKay, and Jainudeen 1971; Estes 1991; Spinage 1994; Su-
kumar 1994; Daniel 1998).
Apprehensive
In situations where an elephant is slightly apprehensive or
ambivalent, unsure of what action to take, it may be ob-
served to engage in a number of confl ict behaviors. For
instance, an individual may stand, listening or observing,
while twisting the tip of its trunk back and forth, or Trunk-
Twisting (Poole 1999a). Or an elephant may Foot- Swing,
raising and tentatively swinging its foreleg (less commonly
the hind leg) intermittently (Moss 1988; Poole 1999a). In
contexts where an elephant feels uneasy, or unsure of the
next action, he or she may also engage in Touch- Face, a
self- directed touching of the face, mouth, ear, trunk, tusk,
or temporal gland, apparently for reassurance (see fi g-
ure 8.4; Kühme 1961; Poole 1999a; Kahl and Armstrong
2000). For comfort, a young individual, typically an infant
Following a Mock- Charge, an elephant takes a step in
the direction of an offending object (elephant, other ani-
mal, human being) and at the end of the movement kicks
up a cloud of dust (Kick- Dust) in the direction of its an-
tagonist. The elephant typically stands tall, towering over
its adversary (Kahl and Armstrong 2000). Often when an
aggressor runs after another and then decides to give up the
chase, he or she extends the trunk upward and outward
toward his or her opponent in an apparent sign of frustra-
tion or “last ditch effort” (Jainudeen, McKay, and Eisen-
berg 1972; Kahl and Armstrong 2000). This Reaching- High
is a common display between competing males, particularly
in association with an estrous female (Poole, personal ob-
servation). During a Dueling contest a male may stretch his
trunk toward his rival as they rush toward one another;
the Trunk- Out- Stretched appears to be used by a male as
a form of defensive protection against the force and sharp
tusks of his rival (Poole 1987b, 1987c). It may be a form of
Distant- Frontal- Attitude, described below.
Redirected- Aggression is aggression aimed at an individ-
ual that is irrelevant to the current situation. When a ten-
dency to attack is thwarted, for some reason (e.g., by fear
of the opponent), an elephant may redirect his or her ag-
gression to some other individual or object, such as thrash-
ing bushes, pushing trees (Bush- Bashing), throwing sticks
and / or grass (Throw- Debris), or threatening or attacking
other, lesser elephants, smaller species, or humans in the vi-
cinity (Sanderson 1907; Spinage 1994; Deraniyagala 1955;
Eisenberg and Lockhart 1972; Krishnan 1972; McKay
1973; Ben- Shahar 1999; Lahiri- Choudhury 1999).
Within elephant families, though not between adult
males, escalated aggression (e.g., Pushing, Tusking) is
often followed by post- confl ict Reconciliation. Typically a
third party, such as the matriarch or a close associate of
the aggrieved individual, initiates the reconciliation. She ap-
proaches the confl icting elephants from the side and, while
standing head to head, rumbles while Head- Raising and
Ear- Lifting (see Affi liative, below) and reaching toward the
other with an outstretched trunk in an affi liative gesture
(Poole, personal observation).
Ambivalent
Displacement
Displacement- Grooming and Displacement- Feeding is be-
havior characterized by its apparent inappropriateness or
irrelevance to the situation in which it occurs. For example,
in confl ict situations an individual may begin to throw dust
or grass on him or herself, when such grooming is inap-
propriate (Kühme 1963; Eisenberg and Lockhart 1972;
Figure 8.4 In situations in which an elephant feels uneasy or unsure of the next
action, he or she may also engage in self- directed touching of the face, mouth,
ear, trunk, tusk, or temporal gland, or Touch- Face. This behavior appears to be a
form of reassurance, or displacement behavior.
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114 Joyce H. Poole and Petter Granli
1981; Poole 1982). As part of social learning calves touch
the mouths of older individuals to sample food items (Lee
and Moss 1999).
During many affi liative or socially exciting interactions
(e.g., when greeting, reaffi rming bonds, expressing soli-
darity, as well as following any exciting event—mating,
birth, fi ght, reconciliation), elephants raise their heads as
in fi gure 8.6. Head- Raising is different from the threaten-
ing Head- High, Standing- Tall posture in that the elephant’s
neck appears to be extended upward and outward so that
the head appears separate from the rest of the body. In the
Standing- Tall posture the elephant’s neck is extended up-
ward but not outward (Poole personal observation). In as-
sociation with Head- Raising elephants lift their ears so that
a gap appears between the lower fl ap and the neck, also
or calf, may engage in Trunk- Sucking (Poole, personal ob-
servation; Kahl and Armstrong 2000).
As an elephant experiences rising apprehension or
alarm, it may adopt several postures that may or may not
be observed in combination. The elephant may stand or
move with the lower potion of the Trunk- Curled- Under
and slightly back (Kühme 1961; Poole 1999a). It may hold
the head and Jaw- Tilted- Upward (so that the tusks become
more parallel to the ground) and the ears stiffened (Ear-
Stiffening) and slightly spread. This posture is unlike that
of an aggressive elephant, who looks squarely at its adver-
sary with ears and tusk more perpendicular to the ground
(Poole 1999a). As an apprehensive elephant moves away
it tilts its head to look back over its shoulder and with its
ears slightly spread and jaw tilted up, gives a characteristic
Tilted- Ear- Spread posture (Poole, personal observation).
As apprehension gives way to increasing fearfulness an ele-
phant raises its tail. Tail- Raising is also observed when ele-
phants are highly excited (Moss 1988; Poole 1999a). An
elephant standing or walking with its jaw tilted upward
and its tail raised causes it to have an exaggerated sway-
back. This overstated Back- Bowing posture (Poole, per-
sonal observation) may also be seen in the Estrous- Walk
adopted by receptive females (Moss 1983). A fearful ele-
phant, or one that is highly aroused, opens its Eyes- Wide so
that the whites may be seen (Poole 1996, 2000a). A fearful
elephant may stand or move while exhibiting a combina-
tion of several displays including Tail- Raising, Eyes- Wide,
Jaw- Tilted- Upward, Tilted- Ear- Spread, Trunk- Curled-
Under, and Back- Bowing (Poole, personal observation).
Once an elephant has decided to retreat, it may run quickly,
quietly and smoothly exhibiting Ear- Flattening and Tail-
Raising. This gait is referred to as Panic- Running. If ele-
phants are in a group they bunch closely together (Moss
1988).
Social Integrative
Affi liative
Elephants engage in numerous forms of low intensity and
more exuberant displays of affi liative behavior that func-
tion to reinforce social bonds between family and bond-
group members. An elephant approaching another family
member may rub (Social- Rub) its head or side against the
other individual as in fi gure 8.5 (Lee [1987]; Moss [1988];
Archie, unpublished). In the context of greeting and reassur-
ing calves an individual may reach its trunk toward or into
the mouth of another (Moss [1981]; Lee [1987]; see Caress,
below). This Test- Mouth behavior is also observed between
non- relatives, especially males in context of greeting (Moss
Figure 8.5 In an affi liative gesture, an elephant approaching another family
member may Social- Rub its head or side against the other individual.
Figure 8.6 During many affi liative or socially exciting interactions, elephants
raise their heads and lift their ears. The Head- Raising and Ear- Lifting posture is
most commonly seen when elephants greet infants or, as in this illustration, their
close associates. Note the posture of the female being approached by the long-
tusked matriarch who is vocalizing with her Mouth- Open. The young female
raises her head and lifts her ears while reaching her trunk back toward the ap-
proaching older female.
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Signals, Gestures, and BehaviorofAfrican Elephants 115
Greeting- Ceremony, as with other Bonding- Ceremonies,
is a constellation of displays including: Head- Raising,
Mouth- Opening, Ear- Spreading, Ear- Lifting, Ear- Folding,
Rapid- Ear- Flapping, Tusk- Clicking, Trunk- Twining, Back-
Toward, Spinning, Temporin secretion, Tail- Raising, Uri-
nation, and Defecation (Moss 1981, 1988; Poole 1987b;
Poole et al. 1988; Poole 2000a).
A Male- Male- Greeting has a very different quality from
a greeting between family members. Males typically ap-
proach one another slowly and reach their trunks into one
another’s mouths. This trunk- to- mouth greeting may be ex-
tended to sniffi ng one another’s temporal glands and geni-
tals, presumably to test one another’s sexual state or iden-
tity (Buss, Rasmussen, and Smuts 1976; Rasmussen 1988).
Greetings may lead to gentle sparring and Trunk- Twining
(see Social Play, below). Greeting males may occasionally
rumble to one another, but they do not vocalize with the ex-
citement shown by greeting family members.
Spatial Proximity
Elephants may use several friendly methods of maintaining
proximity or of monitoring how close another individual is.
While they are moving together in a group one individual
may give another a gentle Herding- Push (see fi gure 8.7),
with the apparent purpose of keeping the group moving to-
gether (Archie, unpublished data). While standing or mov-
ing an elephant may use its tail to forcefully swat an ele-
phant behind it to indicate that it should back off (Kahl and
Armstrong 2000). A gentler Tail- Swat is given to feel what
is behind or to ensure that a calf is still there (Kahl, personal
communication; Poole, personal observation). To push an-
other individual (especially a calf) out of the way, an ele-
phant may Kick- Back with its hind legs (Lee [1987]; Kahl
and Armstrong [2000]; Archie, unpublished data).
seen in fi gure 8.6. This Ear- Lifting posture is most com-
monly seen when elephants greet infants (Coo; Poole, chap-
ter 9) or one another (Little- Greeting; Poole, chapter 9).
As individuals become more aroused Head- Raising and
Ear- Lifting is associated with Rapid- Ear- Flapping, and
often, Ear- Folding (see under Threat). This combination
of postures is almost always seen in the context of greeting
or bonding interactions and during other excited, social,
and chorused calling (Moss 1988; Poole et al. 1988; Poole
1998a; Poole, chapter 9). As excitement levels soar ele-
phants rumble, roar, and scream at extremely high sound
pressure levels (Poole et al. 1988). Covered by the trunk,
the mouths of elephants are not very obvious sources of
visual displays, but during excited vocal interactions ele-
phants open their mouths wide (Mouth- Opening), often
holding their trunks in a curled posture, which exposes the
mouth (Poole, personal observation). During these interac-
tions of extreme social arousal, close associates may gather
together (Gathering) in a close cluster, pressing their bod-
ies against one another. As they vocalize loudly they often
turn abruptly toward and away from each other in a rapid
movement that we refer to as Spinning (Moss 1988). Dur-
ing both lower- and higher- intensity greetings, elephants
may reach out to briefl y seize the trunk of a close associ-
ate (Trunk- Grasping; Poole [1982]; Moss [1988]). Between
greeting males the entwining of trunks is always slow and
sensual; whereas between female relatives during a bonding
or greeting ceremony the action is more sudden and dra-
matic, like a sudden clasping of another during an excited
human greeting (Poole, personal observation). Finally, dur-
ing highly social interactions such as a Greeting- Ceremony
(Moss 1988) or during interactions in which an elephant
expresses solidarity with a family member, elephants may
click their tusks together (Tusk- Clicking; Poole, personal
observation).
These more exuberant displays, known collectively as
bonding ceremonies, may occur during greetings, births
(Poole 1999b), calf rescues, matings (Poole 1987b), or any
number of socially arousing events (see Poole, chapter 9,
for more detail) and involve vocal, chemical, tactile, and
visual communication. During a Greeting- Ceremony, for
example, members of a family or bond group may run to
meet one another while emitting loud, modulated, throaty
rumbles, trumpets, roars, and screams (Moss 1988; Poole
et al. 1988; Poole, chapter 9). The elephants raise their
heads and lift and spread their ears, secreting Temporin pro-
fusely. As they meet they fl ap their lifted ears rapidly, while
rumbling, trumpeting, screaming, and roaring. Rubbing
against one another they stand in parallel and while hold-
ing their heads high, they click tusks together and entwine
trunks. As the greeting continues they may back into one an-
other and spin around, while urinating and defecating. The Figure 8.7 Close associates use a gentle Herding- Push as a friendly way of
keeping the group moving along together.
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116 Joyce H. Poole and Petter Granli
aggressive situations, or to initiate a coalition (Poole, per-
sonal observation).
Mother- Offspring
Suckling
In attempting to gain access to the breast, an infant or calf
walks parallel to its mother, either pushing against her legs
or touching the trunk onto or near to her breast. This beg-
ging or Solicit- Suckling behavior is often accompanied
by a Suckle- Rumble or other begging calls (Lee and Moss
[1986]; Lee [1987]; Langbauer [2000]; Poole, chapter 9).
A mother can reject such solicitations, or Reject- Suck, by
blocking access to her breast with her leg or by walking
away. If the calf is already suckling, the female can termi-
nate nipple contact by stepping forward and pulling the
nipple from the calf’s mouth; she can more aggressively
bump her elbow into the suckling calf to force it off, and
she can slap or push the calf off the nipple (Lee 1986,
1987). Rejections from suckling are often accompanied
by the calf giving protest cries or loud roars (Poole,
chapter 9).
A female suckling an infant, or a juvenile female “pre-
tending” to suckle an infant often holds very still with the
Ears- Lifted in an attentive posture and her eyes opening
and closing or Eye- Blinking (see General Attentive, below)
in what we term a Suckle- Stance (Poole, personal observa-
tion; fi gure 8.9).
Movement Initiation and Leadership
Within families the coordination and initiation of move-
ment is a vital component of group living (Mutinda, Poole,
and Moss, chapter 16). An individual may initiate a move-
ment to a new habitat by adopting a Let’s- Go- Stance,
standing on the periphery of the group, purposefully fac-
ing in the direction she wishes to travel, waiting, and rum-
bling “Let’s go” every minute or so (Poole et al. [1988];
Poole, chapter 9; as shown in fi gure 8.8). In between calling
the elephant waits. Waiting behavior may also be observed
when part of a family is lagging behind. One or more in-
dividuals purposefully wait for the others to either catch
up or to initiate a procession again; contextually this pos-
ture is obvious as the waiting individual is attentive (rather
than resting), often glancing back over her shoulder, and
monitoring the other’s activity with her trunk but other-
wise showing a lack of activity (Moss 1988). The individ-
uals begin to walk again as soon as the individuals they
are waiting for approach. Elephants may indicate to one
another a change of activity or direction by fl apping the
ear (usually one more than the other) moderately loudly
against the neck and shoulder at the same time as the ele-
phant tilts its head upward and to one side, causing the ear
to slide against the shoulder. The Ear- Flap- Slide causes the
ears (or ear) to make a rasping sound as they rub downward
against the shoulders. This signal is especially used by adult
females, but may sometimes be made by adult males and
juveniles of either sex (Moss 1988). A loud Ear- Slap against
the shoulders, typically by a dominant female, appears to be
used to call attention to her location and activity. Dominant
females also use this signal to announce “status” in slightly
Figure 8.8 An individual initiating a movement to a new habitat adopts a Let’s-
Go- Stance and rumbles, “Let’s go” every minute or so. The calling female adopts
a particular posture: she stands on the periphery of the group, sometimes lifting
and swinging a foot (Foot- Swing), and purposefully faces in the direction she
wishes to travel. Her persistent calling attracts the attention of others who may
slowly move to join her.
Figure 8.9 A female suckling an infant (Suckle- Stance) typically holds very still,
Ear- Lifting in an attentive posture.
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Signals, Gestures, and BehaviorofAfrican Elephants 117
is dangerous or potentially dangerous, whether caused by
other elephants, predators (including people), or strange
sounds, smells, or events. Bunching may precede a Full-
Retreat, Panic- Running, or a Group- Charge. Following a
threat to a predator, an individual or group of elephants
may do an about- face and make a rapid and Full- Retreat
from danger. A large adult female places herself several me-
ters behind a bunched group of elephants in Full- Retreat.
This Rear- Guarding female carries her head very high while
looking back over fi rst one shoulder and then the other in
a Look- Back. While this posture indicates an alarmed ele-
phant, the wrong move by the human observer or predator
may provoke an attack (Douglas- Hamilton 1972).
Sexual
Advertisement / Attraction
Among males, the postures associated with musth function
in both male- male competition and aggression as well as
sexual advertisement. For example, musth males carry their
heads well above their shoulders with exaggerated rigidity,
tuck in their jaw, and stiffen their ears. These Head- High,
Chin- In, Ears- Tense postures are elements of a dominant
musth male’s swagger, or Musth- Walk, and characterize all
of their behavior and interactions (Poole and Moss 1981;
Poole 1987a; Kahl and Armstrong 2002; Payne 2003). A
feature of musth is the continual Urine- Dribbling from a
retracted and sheathed penis. The fl ow may vary from an in-
termittent drip to a gushing stream, particularly if the male
is threatening another (Poole 1987a). Males may lose up to
Reassurance and Protection
Using the trunk to touch, or Caress, another in a protective,
reassuring, or comforting manner is a common behavior in
an elephant family. While adult females or juvenile females
most often direct caresses toward infants and calves, adults
and juveniles may also Caress one another. Caresses come
in many forms: wrapping a trunk over the back and around
the belly of a calf or over calf’s shoulder and under its neck,
often touching its mouth; reaching out to touch the genitals,
temporal glands, face, legs, mouth, or trunk of another in-
dividual (Lee [1987]; Poole, personal observation). A gentle
rumble often accompanies a Caress.
Adult and juvenile females are often observed to use the
body, feet, trunk, or tail to shepherd, check for the presence
of, gently guide, or assist an infant or calf to safety. This
Shepherding action may also be used to gauge the proxim-
ity of an elephant behind (Lee [1987]; Kahl and Armstrong
[2000]; Poole, personal observation; Archie, personal com-
munication).
A series of more physically active contacts use the feet or
trunk to retrieve young infants when they are lying down
and potentially in danger of being left behind. For example,
a front or hind foot lifts a calf onto its feet; trunk or trunk
and tusks together pulls or pushes a calf onto its feet, over
a log, or out of a mud wallow. Mothers and allomothers
utilize these behaviors to maintain close contact with and
assist infants (Lee [1987]; Poole, personal observation). Ele-
phants may also use the hind feet, tusks, or trunk to attempt
to lift injured, sick, dying, and dead elephants. They have
even been observed carrying dying infants (Moss 1992).
When adults perceive a threat, they respond by Bunching,
thus protecting calves in the center of a tight defensive circle
of elephants. Adults face outward on all sides, heads up and
moving from side to side, and vocalizing, with their rumps
to the center, like the spokes to a wheel (Lee, personal com-
munication).
Group Defense
When a group of elephants is under threat, they respond
rst by Freezing (see General Attentive, below) and then by
clustering or Bunching together (usually with the youngest
animals in the center) so that the diameter of the group de-
creases (Douglas- Hamilton 1972; Moss 1988; Langbauer
et al. 1989; McComb et al. 2000; see fi gure 8.10). This reac-
tion is different from the Gathering together of family mem-
bers seen during bonding, which also results in a smaller
group diameter. In the former tight group elephants exhibit
a combination of Apprehensive and / or Aggressive postures.
Bunching may be observed in response to any situation that
Figure 8.10 When a group of elephants detects a threat, they respond by clus-
tering or Bunching together (with younger individuals protected in the center).
In this tight group they may exhibit a combination of fearful and / or aggressive
postures as they decide how to respond. The elephants in this illustration wait
apprehensively (note trunks curled under and jaws slightly raised) for the arrival
of the family matriarch before departing.
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118 Joyce H. Poole and Petter Granli
and Armstrong 2002) or lifting the head up and, with open
mouth, reaching with curled trunk to rub across the tem-
poral gland and forehead (Trunk- to- Head; Poole [1987a]).
Musth males may drag, or Trunk- Drag, the distal portion of
the trunk on the ground, making a rasping sound. This be-
havior leaves a snake- like mark on the ground and is likely
to be related to the Trunk- Bounce seen in Asian elephants
(Kahl and Armstrong 2002). Musth males may Trunk- Drag
as a threat at very close range (Poole, unpublished data).
Occasionally in combination with a Trunk- Curl a musth
male raises and lowers his head, or lifts and swings his head
and trunk with vigor, sometimes in gure- eight movement.
In the most intense form of this Head- Toss an elephant
bends his back legs and lowers hind portion of body, caus-
ing the head and trunk to rise even higher (Poole 1987a;
Kahl and Armstrong 2002). Less exaggerated forms may be
observed in non- musth male and female elephants, particu-
larly in play (Moss, personal communication).
A female or a group of females may respond to the
sound, smell, or physical presence of a musth male with a
Female- Chorus display. The female(s) steps forward with
mouth open rumbling and head lifted and outstretched.
Ear postures include a combination of Ears- Lifted and
Rapid- Ear- Flapping. The trunk is extended down and may
be swung slightly as the head is raised. Temporin streams
down the side of the face. The female(s) typically urinates
for the musth male to test and may Rump- Present to him
(Poole et al. 1988; Poole 1999a).
When a non- musth male elephant urinates he fully ex-
tends his penis and usually thereafter obtains a semi-
erection and sometimes a full erection. Infants and calves
usually obtain a semi- erection when they are suckling and
both young and adult males often obtain semi- and full
erections when they are engaged in sparring activities, es-
pecially when these end in Male- Male- Mounting. In some
cases erection appears to be a form of display, though this
requires further investigation (Poole, personal observation).
Sexual Monitoring
The monitoring of sexual state is composed of a medley
of different behaviors, which includes the testing of tempo-
ral glands, urine, feces, and genitals. During the testing of
genitals an individual reaches its trunk toward the genitals
of another (sniffi ng) for the purpose of assessment of re-
productive state (Rasmussen and Wittemyer 2002). Test-
Genitals may be Male- Female, Male- Male, Female- Male,
or Female- Female. In Male- Female- Test- Genitals a male
reaches out with his trunk and sniffs the genitals of a fe-
male (Jainudeen, Eisenberg, and Tilakeratne 1971; Eisen-
berg, McKay, and Jainudeen 1971; Rasmussen et al. 1982).
In Male- Male- Test- Genitals a male reaches out his trunk
300 l per day, and consequently they drink more than non-
musth males (Poole 1989a). The continual dribbling gives
the inner hind legs a black shiny appearance and may turn
the sheath of the penis a greenish color (Poole and Moss
1981; Hall- Martin 1987; Moss 1988). The urine has a very
pungent odor (Poole and Moss 1981; Poole 1987a; Kahl
and Armstrong 2002). Musth males also secrete from the
temporal glands, creating a dark streak down the side of the
face, mid- way between ear and eye. This Musth- Temporal-
Gland- Secretion may be distinguished from Temporin
by its congealed appearance and strong odor (Poole and
Moss [1981]; Poole [1987a] defi nes it using categories of
1–4). Musth males rub their temporal glands against trees
or the ground (when mud wallowing). Marking may be
so vigorous that the male departs with bark and debris on
the side of his face. Dissected temporal glands have been
found to have pieces of bark embedded deep inside them.
Non- musth elephants also mark but the behavior is more
ritualized among musth males (Poole 1987a; Kahl and
Armstrong 2002). Urine- Dribbling and Musth- Temporal-
Gland- Secretion, while primarily olfactory signals, may also
have a visual signaling effect.
Another characteristic behavioral trait of musth males
is the Ear- Wave, as shown in fi gure 8.11. A musth male
swings the upper portion of the ear stiffl y and vigorously
forward. The speed and forcefulness of the swing causes
the lower, unsupported portion of the pinnae to go even
further forward and fl ap upward. The motion creates a
wave appearance across the ear (Poole 1982; Kahl and
Armstrong 2002). The Ear- Wave may function to waft the
odor of Musth- Temporal- Gland- Secretion toward other
elephants (Poole 1987a). In addition, musth males may be
seen to Trunk- Curl, furling and unfurling the trunk in a
sinuous snakelike motion, often curling it over a tusk (Kahl
Figure 8.11 Ear- Wave: A musth male swings the upper portion of the ear
stiffl y and vigorously forward. The speed and forcefulness of the swing causes
the lower, unsupported portion of the pinnae to go even further forward and
ap upward.
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Signals, Gestures, and BehaviorofAfrican Elephants 119
Sexual Solicitation
Musth males often adopt a non- threatening stance or
Casual- Walk when approaching a group of females, while
in the presence of females, or when moving through a group
testing females. This walking (or standing) with the trunk
draped over a tusk in a relaxed and “casual” manner signals
friendly intent (Moss 1988, 1992).
Females may be observed to Back- Towards a male and
Rump- Present for testing or to solicit mating (Poole 1987b;
Poole and Moss 1989; Kahl and Armstrong 2002).
Courtship
Estrus is characterized by a series of specifi c behaviors that
have been described in detail by Moss (1983) and Poole
(1989b). A female in estrous acts with Wariness in the pres-
ence of males, holding her head high, her eyes opened wide,
with her gaze directed at other elephants. She is intoler-
ant of male approaches (Moss 1983) and avoids them with
an Estrous- Walk, a characteristically long gait in an arc
away from family with her head held high and turned to
one side, giving her body a slight twisted motion as she
walks (Moss 1983). The Estrous- Walk becomes a Chase as
a male(s) pursues an estrous female at a fast walk or shuf-
ing run (Moss 1983; Poole 1982, 1989b). Once a pursu-
ing male has caught up with the female he reaches his trunk
across her back in a Reach- Over, or may place his trunk
along her back, and may attempt to prevent her forward
movement (Kühme 1961; Eisenberg, McKay, and Jainu-
deen 1971; Jainudeen, Eisenberg, and Tilakeratne 1971;
Moss 1983). Females are invariably lighter than males and
can thus outrun them; an experienced female chooses to
allow a male to Reach- Over. Once a male has stopped the
forward movement of a female and begun to mount her,
she may either choose to step forward or stand still. Within
an established consort between a musth male and an es-
trous female, the pair may not engage in either the Chase
or Reach- Over phase. Instead, the male may approach the
female and, prior to Mounting, rest his chin on her back or
begin pushing or Driving her, sometimes roughly, with his
forehead (Eisenberg, McKay, and Jainudeen 1971; Poole,
personal observation; see fi gure 8.12). During Driving the
female attempts to stand her ground (Jainudeen, Eisen-
berg, and Tilakeratne 1971; Moss 1983). Standing by the
female involves actively pushing back against the Driving
male with her legs locked. Mounting occurs when a male
has succeeded in placing his forelegs on the female’s back
as depicted in fi gure 8.13 (Moss 1983). Mounting may
be successful (intromission and ejaculation occur) or un-
successful (Poole 1982; Poole 1989b). A successful mat-
ing largely depends upon whether the female chooses to
to smell another male’s genitals. This behavior may occur
following a meeting between two males in association with
Male- Male- Greeting and also by non- musth males toward
musth males in relation to assessment (Poole 1982). In
Female- Male- Test- Genitals, a female reaches her trunk in
the direction of and touches or almost touches a male’s pe-
nis. Females show most interest in musth males and estrous
females display this behavior very prominently immediately
following a mating (see also Test- Semen, below; Eisen-
berg, McKay, and Jainudeen [1971]; Poole [1982, 1987b]).
Female- Female- Test- Genitals is also observed.
In Test- Urine, elephants may also touch the tip of the
trunk over, on, or in urine or a urine spot ( Jainudeen, Eisen-
berg, and Tilakeratne 1971; Rasmussen et al. 1982; Ras-
mussen, Schmidt, and Daves 1986; Rasmussen 1988; Ras-
mussen and Schulte 1998) for the purpose of assessment of
reproductive state (either estrus or musth; Rasmussen and
Wittemyer [2002]) and individual identifi cation (Rasmussen
and Krishnamurthy 2000). Testing of urine may be Male-
Female, Male- Male, Female- Male, or Female- Female. After
an individual places the tip of the trunk in or near a liq-
uid substance (often urine) the trunk may be curled, raised
vertically, and the trunk tip placed on paired orifi ces lead-
ing to the vomeronasal organ in the roof of the mouth in a
Flehmen response (Rasmussen et al. 1982). The vomero-
nasal organ plays an important role in processing chemi-
cal signals relevant to reproduction (Rasmussen et al. 1982;
Rasmussen et al. 1993; Rasmussen 1998). The Flehmen re-
sponse may be observed in males to female urine, to female
estrous urine, and to male musth urine. Sometimes an indi-
vidual may Place the tip of the trunk over a scent of interest
and immediately fl atten the entire end of the trunk over the
sample; sucking or blowing may ensue (Rasmussen 1998).
Placing the tip of trunk over or on dung, or Test- Dung, is
also commonly observed.
Elephants often reach out with their trunks to smell an-
other’s temporal glands. Among females and calves Test-
Temporal- Glands may be a component of the Caress be-
havior and among adult females within a family, sniffi ng
another’s temporal glands may obtain information about
a companion’s emotional or physical state. Greeting males
often sniff one another’s temporal glands and this behav-
ior is probably for assessing physiological and sexual state
(Poole 1982; Rasmussen and Wittemyer 2002).
Musth males are often observed to persistently follow
the scent of another, actively scenting with trunk by sweep-
ing the ground and / or scanning the air. A Tracking musth
male may pursue rivals or estrous females, sometimes for
many kilometers (Poole, personal observation). Moss de-
scribes a musth male who, with aggressive intent, followed
the tracks of her Land Rover for more than a kilometer
(Moss 1988).
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120 Joyce H. Poole and Petter Granli
to be Consorting (Moss 1983; Poole and Moss 1989; Poole
1989b; Kahl and Armstrong 2002). As a female passes the
peak of her estrous period her guarding male begins to lose
interest in her. During this phase the female is often ob-
served overtly Soliciting- Guarding behavior from her ex-
consort by following him, and when approached or pur-
sued by other males she moves rapidly to stand next to him
(Poole 1989b).
Post- Copulatory
Following a mating, the mated female adopts a Post-
Copulatory Stance, stepping forward with streaming tem-
poral glands and Mouth- Opening, she calls with a powerful
series of rumbles (see Poole, chapter 9). With her head
raised and outstretched, she exhibits Rapid- Ear- Flapping.
The female alternately turns toward the male, and then rap-
idly away, holding her head up high and her ears spread
(probably to ensure long- distance advertisement of avail-
ability) and calls powerfully and repeatedly at initially
regular intervals, but then with increasing intervals and di-
minishing intensity (Poole 1987b; Poole et al. 1988; Poole
1989c). These extremely powerful and characteristic calls
attract the attention of distant males (Poole 1987b; Poole
et al. 1988; Poole and Moss 1989). Between calls, the fe-
male may reach her trunk to touch the male’s penis or his se-
men on the ground, sometimes even fl icking semen onto her
body using a trunk movement similar to dusting (Test- Penis
and Test- Semen; Poole [1982]; Poole [1987b]). The mated
female’s relatives rush to her side and join her in exuberant
calling, displaying, urinating, and defecating in what we re-
fer to as a Mating- Pandemonium. They, too, reach to smell
and / or touch the male’s penis or semen. The calling by the
mated female is distinct from the pandemonium produced
by her relatives.
Male- Male
Young males sometimes exhibit sexual behavior toward
one another. For example, a male may chase another male
with an erection much as he would a female. A male with
an erection may be observed to reach over the back of an-
other male, and even mount him, much as he would with
an estrous female (Male- Male- Chase, Male- Male- Reach-
Over, Male- Male- Mounting). The mounting male’s penis is
sometimes rubbed rapidly—using its own muscle- power—
up and down between the other male’s back legs. Ejacu-
lation sometimes occurs (Kahl and Armstrong, personal
communication). This behavior is most often seen between
younger bulls during a friendly interaction (Moss 1988;
Poole 1996).
Stand or move forward. Guarding behavior is said to occur
when a male (usually in musth) attempts to prevent all rival
males from obtaining access to an estrous female by stay-
ing within 5 to 15 m of the female and threatening all ap-
proaching male rivals (Poole 1982, 1989b). When both the
Guarding male and the estrous female exhibit responsibil-
ity for maintaining proximity to one another, they are said
Figure 8.12 A Consorting pair typically precedes a mating with a bout of Driv-
ing. The guarding musth male approaches a female and begins to push her
forward with his trunk or forehead. A female at the height of estrus will attempt
to stand her ground, by locking her forelegs and actively pushing back into the
weight of the male (Standing).
Figure 8.13 Mounting occurs when a male has succeeded in placing his fore-
legs on the female’s back. A successful mount (in which intromission and ejacula-
tion occur) largely depends upon whether the female chooses to Stand or move
forward.
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Signals, Gestures, and BehaviorofAfrican Elephants 121
a used cement bag, a cushion, paper bags, a fi lm canister,
ip- ops, tins, cans, a piece of cloth), though a palm- frond,
swamp vegetation, or a stick will also do (see fi gure 8.15).
This kind of Object- Play, especially with novel objects, can
last 10 minutes or longer with the elephant totally absorbed.
He or she may alternatively kick back at it, roll it under a
foot, touch it with a hind foot, pick it up, pierce it on a tusk,
bite it, swing it around, put it on its head or back, throw
it away, and fi nally, focus attention on it in silent delibera-
tion—only to retrieve it again and begin to toy with it once
more. Similar behavior may be observed when elephants
come across the bones of their own kind. They commonly
pick up an elephant bone, mouth it, bite it, step on it, roll it
under a foot, wave it about, and toss it. The actions while
toying with bones are very gentle and done in quiet contem-
plation (Douglas- Hamilton 1972; Poole 1996; Moss 1992;
Payne 2003). Kicking back at an object, another individual,
Play
Solicit Play
Many of the aggressive and affi liative signals detailed pre-
viously are found in play (see also Lee and Moss, chapter
14). However, when they are used in play, these signals are
(a) out of context and typically out of sequence and (b) ex-
aggerated in their nature and expression. In playful or ex-
pectant situations, an elephant may pause with the trunk
up in a periscope or S- shape waiting for its partner’s next
move. As two individuals approach each other with in-
tent to spar, one (occasionally both) raises the trunk above
the head and curls the tip toward the other individual in
a Distant- Frontal- Attitude (Kühme 1961, 1963). Except
for context, this display appears very similar in form to
Periscope- Sniff (see General Attentive). Individuals may
adopt several other postures in anticipation of, or when
soliciting, a playful interaction. In Solicit- Play an elephant
stretches its head down and out and looks out over its tusks
at a potential playmate. The trunk may be resting folded on
its tusks or placed in its mouth, the ears resting against its
body (Poole, personal observation). In an invitation to play,
or when an individual is considering an object of play or en-
gaging in play it gently waggles its head from side to side
(Head- Waggle; Poole [1996]; Kahl and Armstrong, per-
sonal communication). To solicit play with younger individ-
uals, an older calf lies down, or gets down on its knees (Lee
1986; Poole 1998b), thereby appearing smaller, and turning
off the threat latent in many play moves. This behavior ap-
pears to attract younger calves who wander over and climb
on top or Climb on (see Social Play, below) the older indi-
vidual. In similar fashion, a larger male may lower himself
down on his knees, Kneel- Down, in order to playfully spar
with a smaller male who has shown signs of being afraid to
participate (Moss 1988; Poole 1996; see fi gure 8.14).
Lone and Object- Play
Young elephants, in particular, engage in numerous forms of
object and lone- locomotor play. An elephant may sit down,
raise its trunk high in the air and then allow it to fl op down
on its head, and repeat these actions over and over (Sitting;
Poole, personal observation). Trunk- Squelching is an often
repeated wrinkling up of the trunk and forcing air through
it to produce a “squelching” sound (Poole, chapter 9). Or
an elephant may suck water into its trunk and then spray it
out while swinging its trunk like a child with a garden hose
(Poole and Granli, personal observation). This appears to
be done for amusement.
Elephants are very curious and are quick to investigate
and play with anything new in their environment. Elephants
are particularly engaged by novel man- made “toys” (e.g.,
Figure 8.14 When soliciting play with younger individuals, an older calf may
attempt to appear smaller by getting down on its knees, Play- Kneel- Down, or
lying down, which stimulates younger animals to clamber on top in a Play- Climb-
Upon. In this photograph the older individual also displays a Play- Tusk- Ground.
Figure 8.15 In Object- Play, an elephant’s curiosity is easily stimulated by a
novel object. The “toy” may be kicked or touched with a hind foot, rolled under
a foot, picked up, pierced by a tusk, bitten, swung around, placed on the head
or back, tossed, and then quietly contemplated.
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122 Joyce H. Poole and Petter Granli
another with the head, ears, and tail raised. Play- Pursuit
partners switch between chasing and being chased (Lee
1986). One individual may grasp the tail (Grasp- Tail) of
another, and can pull on the tail (or other body parts) in a
playful situation (Moss 1988) During periods of play, and
often after a play- chase, young calves and juveniles of both
sexes Play- Mount others irrespective of the other’s sex (Lee
1986; see Lee and Moss, chapter 14).
In a gentle form of play, elephants Play- Social- Rub,
pushing at the body, head, or legs of another, who may be
lying down or playing with another elephant (Lee 1986).
Young calves will also climb on top of another elephant
who is lying down, pushing and shoving against the lower
elephant, or they will lie fully ventrally on the other and
wiggle. This game of Climb- On play may lead to a big pile
of wiggling, kicking out, and squirming elephants (Lee
1986; Moss 1988; Poole 1996). This behavior also can
be a prelude to chases, Sparring, or gentle Trunk- Twining
between two calves (Lee 1986; Moss 1988).
Generally Attentive
Finally, there are a number of postures and gestures that are
components of an elephant’s sensory input. A Sniff- Toward
is the most commonly used form of sniffi ng. The trunk
is held relatively straight and pointed in the direction of
interest. The trunk lifted up in an s- shape, or Periscope- Sniff
(see fi gure 8.16), is used to detect scents carried on the wind
(Rasmussen and Schulte 1998; Poole 1999a) and is particu-
larly used when additional information is required, such as
or simply at the air is a common action during play (Play-
Kick- Back) that may be repeated for long periods, at an
object of interest. While two or more play together, an ele-
phant may guard his “toy” from access by another. Other
youngsters may queue up and appear to wait quietly for
their turn with the toy (Lee 1986; Moss 1988; Poole1996,
1998b; Poole and Granli 2003, 2004).
As play becomes more exuberant an elephant may tusk
the ground and lift up vegetation (Kühme 1961; Poole
1987a, 1996). Play- Tusk- Ground is often a form of lone
play although it can also occur during social play. Exuber-
ant play is usually associated with Play- Trumpets (Poole,
chapter 9), which attract the interest of other elephants and
leads to social play. Highly energized, playful elephants en-
joy running back and forth through long grass or shrubs
(Play- Bush- Bashing) or lifting up and throwing objects
(Play- Throw- Debris). Many of these behaviors—Tusk-
Ground, Bush- Bashing, and Throw- Debris—are observed
in escalated contests between musth males.
However, in play elephants adopt contradictory pos-
tures. They stand or run about in Feigned- Fear, adopt-
ing the Jaw- Tilted- Upwards and Tail- Raising as if highly
alarmed. With Eyes- Wide they stare out over their tusks
at “imaginary enemies,” alternately charging in Bow-
Neck type posture or running away as if fearful (Moss
1988; Poole 1996). As part of this high- spirited play ele-
phants spin or charge and turn about (Pirouette- Run), often
Standing- Tall or Head- Shaking, and Trumpeting. Unlike an
aggressively charging elephant, the running gait of a playful
elephant is loose and fl oppy; the individual shakes a low-
ered head from side to side, allowing its ears to fl ap against
its neck and curling its tail up high. Floppy- Running is as-
sociated with Pulsated- Trumpets (Moss 1988; Poole 1996;
Poole, chapter 9).
Social Play
The best- known type of social play is Sparring, which
occurs predominantly between males. Young males in par-
ticular test one another’s strength by placing their trunks
on one another’s heads and pushing their opponent down
and back. Unlike fi ghting males, the ears are relaxed. Spar-
ring can range from gentle trunk wrestling and pushing in
a playful or greeting context, and more boisterous shov-
ing, to rough and aggressive tusking (Kühme 1961; Poole
1982; Lee 1986; Langbauer 2000). Sparring typically
occurs between males, less frequently between a male and
female, and rarely between two females (Poole, personal ob-
servation). Sparring may be initiated by a bout of Trunk-
Twining, a gentle contact form of play where two elephants
twist their trunks together in a spiral (Lee 1987). It may be
interrupted by a pause in which one individual runs after
Figure 8.16 An elephant’s nose is extraordinarily fl exible and may sniff toward
an object of interest in a variety of positions, which tend to be adopted in spe-
cifi c circumstances. The elephant in the foreground uses a Periscope- Sniff to
detect a scent carried on the wind. This position is frequently adopted when ad-
ditional information is required, such as when meeting strangers. The smallest
elephant uses a J- Sniff, often seen when elephants are monitoring an ongoing
situation. The other two use the Sniff- Toward.
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Signals, Gestures, and BehaviorofAfrican Elephants 123
it simultaneously raises its head and stiffens its ears. Some-
times an elephant may turn its head from side to side to
localize a sound (Poole et al. 1988; Heffner et al. 1882).
Similarly, during Freezing an individual or group of individ-
uals suddenly ceases all movement, holding stock- still, ap-
parently Listening or feeling distant vibrations (Moss 1988;
Langbauer et al. 1989).
African elephants normally stand or move with their eyes
cast down. A direct gaze with Eyes- Open is a component
of many postures. An elephant may look in the direction
of interest with its Eyes- Open, visually attending to objects
(Poole, personal observation). An attentive elephant may
slowly open and close, or blink its eyes. This Eye- Blinking
behavior is perhaps most noticeable during suckling, but
can be combined with apprehensive head tilting, or ob-
served when an elephant is attentive and otherwise motion-
less (Poole, personal observation; Lee, personal observa-
tion). When Monitoring an unfolding situation, particularly
an interaction, an attentive individual holds its ears slightly
lifted and partially extended, and gazes with Eyes- Open or
Eyes- Blinking at the subject of interest. Simultaneously, the
elephant sniffs by pointing its trunk tip in the direction of
interest, the fi ngers of the trunk typically opening and clos-
ing (Poole, personal observation).
In Explore- Touch an elephant uses a foot (particularly
the hind foot) or its trunk to investigate an elephant or ob-
ject. This behavior can be seen in object play and is often as-
sociated with Play- Kick- Back, or as a form of Shepherding,
but is perhaps most clear and prolonged when an elephant
investigates a dead elephant or elephant bones when it may
touch or stroke the body or bones of the deceased animal
(Douglas- Hamilton 1972; Moss 1988; Moss 1992; Poole
1996; Payne 2003; McComb, Baker, and Moss 2006).
Death
A description of elephant behavior would be incomplete
without mentioning their behavior around injured, sick,
dying and dead elephants, and their remains. Similar to
the Retrieving behavior observed in the care of infants, ele-
phants may use the tusks, trunk, or feet to attempt to lift
and even carry sick, dying, or dead elephants (Douglas-
Hamilton 1972; Moss 1992; Poole 1996; Payne 2003;
Douglas- Hamilton et al. 2006). Males may attempt Mount-
ing a dead elephant. Elephants have been observed to feed
those who are not able to use their own trunks to eat (Mo-
ses Kofi Sam, personal communication) and to attempt to
feed elephants who have died (Croze, cited in Moss [1982]).
They are also known to collect vegetation and dirt with the
trunk, feet, or tusks and use it to cover a dead elephant or
dead human (Moss 1992; Poole 1996). When an elephant
when meeting strangers or danger (Poole, personal obser-
vation). In the Hovering- Sniff, the trunk is held suspended
over a particular scent on the ground or object of interest
(Rasmussen and Schulte 1998; Poole 1999a). The shape of
the trunk may take many forms and may be surreptitious, as
in the J- Sniff, where the trunk tip is curved slightly back as
if the elephant is attempting to appear not overly interested.
Younger males in the company of a musth male and an es-
trous female often employ this surreptitious form of sniff-
ing, as if not to draw undue attention to themselves. When
an elephant is curious about an object or scent, but dares
not approach more closely, it reaches out with a full exten-
sion of the trunk in a Horizontal- Sniff to inspect the item
(Poole, personal observation; Kahl and Armstrong 2000).
During Tracking, an elephant’s trunk moves regularly back
and forth following a scent trail along the ground as the
(usually) male walks purposefully and persistently in pro-
longed pursuit of another individual (Poole 1999a). This
behavior can also be seen when an elephant tracks a person.
Elephants are capable of discriminating between the
voices of elephant individuals over distances of up to 2 km
(McComb et al. 2003); such auditory recognition is associ-
ated with listening behavior. Listening behavior is also ob-
served when an elephant is aware of other socially interest-
ing sounds or those associated with potential danger. An
individual rarely stands stock- still except when Listening
or resting; usually some part of the body, ears, trunk, or tail
is in motion. A resting elephant relaxes its head and ears,
allowing its head to hang below its shoulders and its ears
to fl op forward. A Listening elephant stands with its head
raised and its ears lifted and slightly extended at an angle
of at least 45 degrees (see fi gure 8.17). The body and ex-
tremities of a Listening elephant suddenly cease moving, as
Figure 8.17 A Listening elephant stands with its head raised and its ears lifted
and slightly extended at an angle of about 45 degrees or more. The body and
extremities of a listening elephant suddenly cease moving, as it simultaneously
raises its head and stiffens its ears. Sometimes an elephant may turn its head
from side to side to localize a sound.
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124 Joyce H. Poole and Petter Granli
Some behaviors are visually dramatic and powerful, even
terrifying to the casual observer. Other gestures are exqui-
sitely understated, requiring a highly trained eye to dis-
cern the subtle shift in movement and posture. This chap-
ter has been an attempt to introduce others to many of the
behaviors we use in our quest to understand and describe
the social, emotional, and cognitive world of African ele-
phants. These behaviors are, in effect, our “tool box.” Over
many years, we have learned that certain relationships (e.g.,
mother- infant, mother- daughter, male competitors, sexual
partners, friends, foes) are characterized by specifi c behav-
ioral patterns made up of explicit components. Armed with
this knowledge, we feel confi dent in interpreting elephant
behavior, interactions, and relationships wherever we fi nd
them. We hope that through this chapter we have been able
to impart some of our knowledge for others to use toward
the furthering of scientifi c understanding and improving
the conservation and welfare of elephants wherever they
may be.
has been responsible for the death of a person or another
animal they have been observed to vigorously fl atten the
ground (Trample- Ground) around the animal (Poole 1996).
Similar behavior is seen when a baby is born (Poole 1996).
Finally, elephants may engage in Body- Guarding, standing
over the body of a dead elephant or person and protecting
it from the approaches of predators or other elephants, and
threatening approaching individuals (Croze, cited in Moss
1982; Douglas- Hamilton 1972; Poole 1996; Payne 2003).
As mentioned before, elephants typically also investigate
the remains of elephants sniffi ng with the trunk, using the
feet and trunk to Explore- Touch, lifting, carrying, and play-
ing with the bones or considering them in quiet refl ection
(Douglas- Hamilton 1972; Moss 1988, 1992; Poole 1996;
Payne 2003; McComb, Baker, and Moss 2006).
Conclusions
The visual and tactile signals, gestures, and behaviors of
African elephants are extremely intricate and highly varied.
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... Elephant reactions to sample presentation were documented by scan sampling with an ethogram derived from Poole and Granli [23,24], which consists of 27 behaviors (see Table 2). These behaviors were aggregated into four higher-level behavioral categories: neutral, excitement, mental processing, and sample examination. ...
... Even though this study was performed only on four elephants, data suggest that elephants can recognize the scent of their relatives after up to 12 years of separation. It thereby also demonstrates the intense social bond between elephants [11,12,23]. Even after 12 years of absence, the scent of a relative caused reactions of excitement (see also Video S1). ...
... The elephants studied here exhibited all behavioral categories associated with agitation and connected the scent to their relative, whereas only minor interest and agitation but major neutral behavior was shown during the non-kin sample presentation. The excitement behavior ( Table 2) that was observed during the trials can be assumed as a positive association, as indicated by the expressed rumbling noises and the ear flapping, which are both classified as positive agitation [23,24], and their repeated examination of the sample. Hence, the sample presentation of the absent kin led to a positive reaction of possible emotions, which means that behavioral and bonding concepts by the elephants with their family members are still present ex situ and do not get lost in the zoo environment [19,20]. ...
Article
Full-text available
Simple Summary African elephants are known for their long memory; this is also valid for their olfactory sense and their ability to discriminate scents. This feature is highly important for these mammals to maintain their family bonds and to differentiate between familiar and unfamiliar individuals. Thus far, scientific data only testify to an olfactory memory of up to one year for African elephants. This study investigated the long-term olfactory memory of two mother-daughter pairs that were separated for 2 and 12 years, respectively. Results showed that all four elephants were able to recognize their separated relatives just by the scent of feces, thereby giving the empirical implication of olfactory memory in African elephants of up to 12 years. Abstract African elephants are capable of discriminating scents up to a single changed molecule and show the largest reported repertoire of olfactory receptor genes. Olfaction plays an important role in family bonding. However, to the best of our knowledge, no empirical data exist on their ability to remember familiar scents long-term. In an ethological experiment, two mother-daughter pairs were presented with feces of absent kin, absent non-kin, and present non-kin. Video recordings showed reactions of elephants recognizing kin after long-term separation but only minor reactions to non-kin. Results give the empirical implication that elephants have an olfactory memory longer than 1 year and up to 12 years and can distinguish between kin and non-kin just by scent. These findings confirm the significance of scent for family bonds in African elephants.
... In addition, musth status was evaluated based on the presence of musth-specific traits, such as active urine dribbling, urine staining on the back legs, temporal gland swelling, and secretions [11,12,61,72]. Behaviors known to be associated with musth-such as ear waving, trunk curling, trunk dragging, and the musth walk-were also recorded [12,61,72,73]. Elephants were considered in musth if a suite of these visible signs were displayed during behavioral observations. ...
... A score of 0 (0%) was assigned if target males exhibited no apparent reaction to the call and continued on their directional trajectory out of the clearing. By contrast, a score of 1 (100%) represented the greatest response intensity and was characterized by males tracking [61,73] and approaching the speaker. If males reacted to the call by immediately freezing, listening, and/or orienting their entire body once towards the direction of the sound (vigilant and attentive behaviors, see Table 2) [61,73,74], they were considered to have a response intensity of 25% (score = 0.25). ...
... By contrast, a score of 1 (100%) represented the greatest response intensity and was characterized by males tracking [61,73] and approaching the speaker. If males reacted to the call by immediately freezing, listening, and/or orienting their entire body once towards the direction of the sound (vigilant and attentive behaviors, see Table 2) [61,73,74], they were considered to have a response intensity of 25% (score = 0.25). This score represents the minimum reaction to the estrous call, rather than to another potential sound or environmental stimulus. ...
Article
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Driven by reproductive motives, male African elephants (Loxodonta africana) in musth often expand their home ranges to locate estrous females. This extended range, coupled with heightened aggression often observed in musth males, can be particularly problematic in regions where human-modified landscapes and elephant territories increasingly overlap. Several mitigation tools have been tested to resolve a wide range of human–elephant conflicts with varying degrees of success due to geographical disparities and habituation. We present findings on the potential application of estrous call playbacks in manipulating the behavior and movement of male elephants non-invasively, particularly mature musth adults and younger post-dispersal males, in Etosha National Park. Estrous vocalizations were presented across 26 experimental trials to mature musth adults (n = 5), mature non-musth adults (n = 6), and non-musth males belonging to younger, post-dispersal age classes (n = 8), with behavioral responses scored on a gradient scale from 0–1. Both mature musth adults and younger non-musth elephants were significantly more likely to respond with the highest intensity by approaching the acoustic source compared to mature non-musth adults that avoided the call. However, younger males tested in the presence of an older, higher-ranking male tended to react with a lower intensity than those tested alone. This result likely demonstrates the influence of social hierarchy and associations on male elephant behavior. We also observed a significant increase in physiological response, measured by defecation rate, across all male groups in response to the estrous call playbacks. Our findings suggest that using estrous calls as acoustic deterrents may effectively and non-invasively aid in reducing tension at the human–elephant interface, depending on the age, social context, and reproductive status of the male elephant.
... While the increased probability of performing aggressive behaviours to non-elephant targets when in higher-risk social contexts may represent responses to targets actually perceived as threatening by elephants with a heightened sensitivity, this aggression may alternatively or additionally be a form of re-directed or displaced aggression linked to an acute stress response induced by a perceived threatful social condition [39,55]. Indeed, aggression to non-elephant targets often appeared not to be a true anti-predator defence because it was directed at non-threatening objects or bystanders (for example, bashing of vegetation, charging of birds or smaller ungulates) or had no obvious target (target was unidentifiable, see electronic supplementary material, figure S3). ...
... Ethogram of behaviours recorded during focal follows and their categorization for analysis in the current study[39,40].conspecific aggressionAggressive behaviours relating to dominance assertion and gaining access to resources, as well as potentially redirected aggression including 'advancing towards', 'spreading ears', 'holding head high', 'ear folding', 'head shakes', among other behaviours (electronic supplementary material, note S3 for full list of behaviours and detailed descriptions) directed by the focal subject towards conspecifics. aggression directed to non- ...
Article
Full-text available
Males in many large mammal species spend a considerable portion of their lives in all-male groups segregated from females. In long-lived species, these all-male groups may contain individuals of vastly different ages, providing the possibility that behaviours such as aggression vary with the age demographic of the social environment, as well as an individual's own age. Here, we explore social factors affecting aggression and fear behaviours in non-musth male African elephants ( Loxodonta africana ) aggregating in an all-male area. Adolescent males had greater probabilities of directing aggressive and fearful behaviours to non-elephant targets when alone compared to when with other males. All males, regardless of age, were less aggressive towards non-elephant targets (e.g. vehicles and non-elephant animals) when larger numbers of males from the oldest age cohort were present. The presence of older males did not influence the probability that other males were aggressive to conspecifics or expressed fearful behaviours towards non-elephant targets. Older bulls may police aggression directed towards non-elephant targets or may lower elephants’ perception of their current threat level. Our results suggest male elephants may pose an enhanced threat to humans and livestock when adolescents are socially isolated, and when fewer older bulls are nearby.
... Social linkages and the maintenance of contact between individuals with means to identify the caller and its group affiliation have been noted in sperm whales and their use of overlapping, duet-like sequences of coda calls [100,171] and the whistles of dolphins [142,172]. Contact calls to facilitate group aggregation and reunion have been noted in walruses (e.g., [101]); bats (e.g., [173][174][175]), and elephants (e.g., [176][177][178][179]). Group or pack calls described as 'rallying calls' have been noted for primates, wolves, hyenas, coyotes, and lions [180]. ...
Article
Full-text available
The behavioural, physiological, and energetic repercussions for wildlife that result from changes in their soundscapes are increasingly being realized. To understand the effects of changing acoustic landscapes, we first must establish the importance of the acoustic sense for species to transfer information between the environment, con- and heterospecifics, and a receiver, and the functional role of calling in behaviours such as foraging, navigation, mate attraction, and weaning. This review begins with a discussion of the use of calling and the acquisition of the vocal repertoire, before providing examples from multiple taxa on the functional applications of signals and communication. The acoustic sensory mode adds to, if not being inherent in, many critical life history stages over a range of species. The potential effects on an animal resulting from a change in its perceived soundscape and disturbance on its acoustics use is outlined. This can then be used to consider the implications of an altered acoustic niche or active space in the success and survival of an individual or species. Furthermore, we discuss briefly metrics that could be used to understand the implications of these changes, or could be used to guide mitigation action to lessen the impact.
... For example, elephants live in complex social groups where adult females cooperate in group defence, o spring care, and resource acquisition, and were shown to possess a rich multimodal communication system that includes vocal, visual, tactile, and chemical signals that can encompass speci c meanings and intents shared with other members of the herd (Kahl & Armstrong, 2000;Langbauer, 2000;Poole & Granli, 2011;Soltis et al., 2005aSoltis et al., , 2005bStoeger, 2021;Wierucka et al., 2021). This complex signal system mediates the intricate teamwork displayed by members of an elephant family, including day-to-day decision-making such as deciding when and where to go or how to respond to predators (Jacobson & Plotnik, 2020). ...
Chapter
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... Elephants display an impressive capacity for short-and long-range communication, and rely on a rich multimodal sensory system that includes vocal, visual, tactile, and chemical signals. Their acoustic communication system includes an extensive vocal repertoire (e.g., trumpets, roars, low-frequency rumbles), which encompasses specific functions and intents shared with other members of the herd (45,128,129,134,135,138,178). For example, elephants in Kenya have different alarm calls for humans and for bees, which elicit different responses (134,136). ...
Article
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Humans are unique in their sophisticated culture and societal structures, their complex languages, and their extensive tool use. According to the human self-domestication hypothesis, this unique set of traits may be the result of an evolutionary process of self-induced domestication, in which humans evolved to be less aggressive and more cooperative. However, the only other species that has been argued to be self-domesticated besides humans so far is bonobos, resulting in a narrow scope for investigating this theory limited to the primate order. Here, we propose an animal model for studying self-domestication: the elephant. First, we support our hypothesis with an extensive cross-species comparison, which suggests that elephants indeed exhibit many of the features associated with self-domestication (e.g., reduced aggression, increased prosociality, extended juvenile period, increased playfulness, socially regulated cortisol levels, and complex vocal behavior). Next, we present genetic evidence to reinforce our proposal, showing that genes positively selected in elephants are enriched in pathways associated with domestication traits and include several candidate genes previously associated with domestication. We also discuss several explanations for what may have triggered a self-domestication process in the elephant lineage. Our findings support the idea that elephants, like humans and bonobos, may be self-domesticated. Since the most recent common ancestor of humans and elephants is likely the most recent common ancestor of all placental mammals, our findings have important implications for convergent evolution beyond the primate taxa, and constitute an important advance toward understanding how and why self-domestication shaped humans' unique cultural niche.
... However, like the other elephants, N'Dala still managed to grasp all the food items. These results confirms the idea that vision, one of elephants' less-used senses (O'Connell-Rodwell, 2007;Poole & Granli, 2011;Plotnik et al., 2013), is not necessary for grasping food items with the trunk. ...
Article
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Background During reach-to-grasp movements, the human hand is preshaped depending on the properties of the object. Preshaping may result from learning, morphology, or motor control variability and can confer a selective advantage on that individual or species. This preshaping ability is known in several mammals ( i.e., primates, carnivores and rodents). However, apart from the tongue preshaping of lizards and chameleons, little is known about preshaping of other grasping appendages. In particular, the elephant trunk, a muscular hydrostat, has impressive grasping skills and thus is commonly called a hand. Data on elephant trunk grasping strategies are scarce, and nothing is known about whether elephants preshape their trunk tip according to the properties of their food. Methods To determine the influence of food sizes and shapes on the form of the trunk tip, we investigated the morphology of the distal part of the trunk during grasping movements. The influence of food item form on trunk tip shape was quantified in six female African savannah elephants ( Loxodonta africana ). Three food item types were presented to the elephants (elongated, flat, and cubic), as well as three different sizes of cubic items. A total of 107 ± 10 grips per individual were video recorded, and the related trunk tip shapes were recorded with a 2D geometric morphometric approach. Results Half of the individuals adjusted the shape of the distal part of their trunk according to the object type. Of the three elephants that did not preshape their trunk tip, one was blind and another was subadult. Discussion and perspectives We found that elephants preshaped their trunk tip, similar to the preshaping of other species’ hands or paws during reach-to-grasp movements. This preshaping may be influenced by visual feedback and individual learning. To confirm these results, this study could be replicated with a larger sample of elephants.
Article
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South Africa has many fenced reserves harbouring small to medium sized populations of African elephant ( Loxodonta africana ), most of which have been translocated. Elephants on fenced reserves may be exposed to various management interventions and practices (translocation, hunting, darting, high tourism impact, contraception programs, disruption due to infrastructure maintenance, etc.). These factors may impact the welfare of elephants. Poor elephant welfare may have serious consequences such as increased inter- and intra-species aggression that could result in fatalities. This is the first study to attempt to define behavioural and physiological welfare parameters for free-ranging elephants on small to medium sized reserves. The eight study sites incorporated reserves with different social structure combinations, elephant life-histories, reserve sizes, habitat, management, and tourism intensity. Data collection consisted of behavioural observations (10-minute videos) as well as faecal samples. By incorporating both behavioural and physiological (faecal glucocorticoid metabolite (fGCM) concentration) parameters, we aimed to investigate whether the two parameters showed similar trends. Five behavioural categories were identified (Arousal, Assessing, Ambivalent, Ambivalent/ Body care, and Frustrated behaviour), with various detailed behaviours demonstrated by the elephants that may indicate the influence of anthropogenic disturbance and possibly impact on animal welfare. The study showed significant differences between the selected detailed behaviours, behavioural categories and fGCM concentrations of elephants across the eight reserves. History seemed to be a decisive factor, as reserves with predominantly ex-captive elephants showed higher frequencies of certain behaviours as well as higher fGCM concentrations. Age, sex, reserve size and season were also found to contribute to our defined welfare indices and fGCM concentrations. This indicates that behavioural parameters, indicative of certain behavioural states, are valuable indicators of welfare, as supported by the physiological response of the elephants. The results also highlight the importance of taking multiple specified behaviours from a category into consideration when evaluating the welfare of elephants, to account for individual variation.
Thesis
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As an endangered mammal, novel research into the African elephant’s ethochemistry will assist ethologists, conservation biologists and wildlife managers in understanding odour signals in elephant exudates, and how they impact elephant behaviour in captivity and in the wild. Analytical methods combining genetics, statistics with SPME GC_MS and innovative metabolomics visualisation software are a useful first step in unlocking odours salient to elephant reproduction, breeding and behaviour. Mammals, such as the African elephant, which live in sophisticated, fission-fusion, social systems marked by linear dominance hierarchies and ritualized greeting ceremonies, require an effective means of communication in order to maintain social cohesion. A number of acoustic, olfactory and visual signals have been described. Within a group context, individuals need to discriminate between members of their group and foreign conspecifics in order to avoid inbreeding and to identify recipients of nepotistic behaviour. Such olfactory group/herd/clan/colony identity signals have been described in a number of mammals such as beavers, bats, naked mole rats, rabbits, fur seals, lemurs and hyena but are lacking in the iconic African elephant. It is to date not clear what fixed and variable information is encoded in African elephant urine, temporal gland secretions (TGS), genital and buccal secretions, and whether an olfactory signal for herd membership exists, although scientists have long expected this to be the case. Specifically, African elephant TGS, buccal and genital secretions in adult females have not been described in a free-ranging population. The overall objective of this research was to augment scientific understanding of African elephant olfactory behaviour in a multi-disciplinary way, combining behavioural observation and experimental trials of African elephants in sanctuaries in South Africa, with genetic and chemical data sampled from a free-ranging African elephant population during a translocation event in Malawi. The research first assesses olfactory acuity and scent discrimination ability in African elephants through novel behavioural bioassays. Subsequently, this study examines whether chemical signals in urine, TGS, buccal and genital secretions encode for age, sex and identity, and explores whether odour is correlated with genetic relatedness. For the first time, the chemistry in TGS, buccal and genital secretions is characterised in wild African elephant adult females (n=40). Elephants were found to be highly adept at scent-discriminating between humans, and scent- tracking a target human across various substrates, including distractor trails. Results suggest that elephants may utilise olfaction to discriminate between individual conspecifics. This was tested using behavioural bioassays, which concluded that African elephants can discriminate between unfamiliar and familiar conspecifics from both urine and dung. Urine elicited the most interest, and was used for further detailed analysis using the comprehensive metabolomics tool XCMS online. TGS, buccal and genital secretions from wild-sampled elephants (n=113) were found to encode individual identity in African elephants. TGS and buccal secretions encoded for age, and TGS and genital secretions tended towards significance for sex. Genetic relatedness was not correlated with odour in TGS, buccal and genital secretions at the herd level. However in all three odours, Similarity Percentage Analysis (SIMPER) identified “herd” as a highly significant factor in explaining chemical differences between herds. This suggests that group odour in African elephants is not correlated to genetic relatedness at the population, herd and individual level, and allows for the possibility that bacteria may be involved in group odour. The biomarkers alkan-2-ones and alkan-2-ols previously published in association with elephant male urinary microbial communities, were identified in adult female buccal and labial secretions. SIMPER analyses further extracted a number of short-chain fatty acids significant to chemical differences between groups (notably acetic acid, pentanoic acid, hexanoic acid, decanoic acid, dodecanoic acid and tetradecanoic acid). These and other compounds have been identified in similar studies as key volatile compounds from bacterial fermentation in clan-living hyenas and colonially-living meerkats, further supporting this possibility. The frequent affiliative social behaviour of African elephants is posited as a likely mechanism for bacterial transmission. Conservation objectives can be supported by an improved understanding of how chemical communication in social mammals like elephants influences mate choice, establishing social relationships, courting, mating and the use of scent for establishing territories. In African elephants, an improved understanding of elephant semiochemistry and eliciting behaviour could shed light on scent manipulation for elephant translocation and reintroductions, captive breeding, welfare and enrichment protocols, human elephant conflict (HEC), elephant reintroductions, social behaviour and habitat selection.
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The social complexity hypothesis (SCH) for communication states that the range and frequency of social interactions drive the evolution of complex communication systems. Surprisingly, few studies have empirically tested the SHC for vocal communication systems. Filling this gap is important because a co-evolutionary runaway process between social and vocal complexity may have shaped the most intricate communication system, human language. We here propose the African elephant Loxodonta spec. as an excellent study system to investigate the relationships between social and vocal complexity. We review how the distinct differences in social complexity between the two species of African elephants, the forest elephant L. cyclotis and the savanna elephant L. africana, relate to repertoire size and structure, as well as complex communication skills in the two species, such as call combination or intentional formant modulation including the trunk. Our findings suggest that Loxodonta may contradict the SCH, as well as other factors put forth to explain patterns of vocal complexity across species. We propose that life history traits, a factor that has gained little attention as a driver of vocal complexity, and the extensive parental care associated with a uniquely low and slow reproductive rate, may have led to the emergence of pronounced vocal complexity in the forest elephant despite their less complex social system compared to the savanna elephant. Conclusions must be drawn cautiously, however. A better understanding of vocal complexity in the genus Loxodonta will depend on continuing advancements in remote data collection technologies to overcome the challenges of observing forest elephants in their dense rainforest habitat, as well as the availability of directly comparable data and methods, quantifying both structural and contextual variability in the production of rumbles and other vocalizations in both species of African elephants.
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