Elephant sociality and complexity: The scientific evidence

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ELEPHANTS AND ETHICS
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Edited by
CHRISTEN WEMMER AND CATHERINE A. CHRISTEN
ELEPHANTS
AND ETHICS
TOWARD A MORALITY OF COEXISTENCE
Foreword by John Seidensticker
THE JOHNS HOPKINS UNIVERSITY PRESS
BALTIMORE
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Elephants and ethics: toward a morality of
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Seidensticker.
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1. Elephants—Effect of human beings on—Moral
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and ethical aspects. I. Wemmer, Christen M.
II. Christen, Catherine A. (Catherine Ann)
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Chapter Opening Art Credits
1: Jordana Meyer
2: Smithsonian Institution Archives,
SIA2008-0924
3: Donna Nissani
4: Petter Granli / elephantvoices.org
5: Courtesy John Edwards, London
6: Jessie Cohen, Smithsonian’s National Zoo
7: Janine L. Brown
8: Disney’s Animal Kingdom
9: Mehgan Murphy, Smithsonian’s National Zoo
10: Smithsonian Institution Archives,
SIA2008-0922
11: Feld Entertainment, Inc.
12: Smithsonian Institution Archives,
SIA2008-0926
13: Disney’s Animal Kingdom
14: Jessie Cohen, Smithsonian’s National Zoo
15: Disney’s Animal Kingdom
16: Prithiviraj Fernando
17: Petter Granli / elephantvoices.org
18: Hank Hammatt
19: Petter Granli / elephantvoices.org
20: Jordana Meyer
21: Stephen Blake, Wildlife Conservation Society
22: Joyce Poole / elephantvoices.org
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V. Krishnamurthy
(1927–2002)
This volume is dedicated to the
memory of Dr. V. Krishnamurthy,
a remarkable Forest Veterinary Surgeon
of India’s Tamil Nadu Forest Department.
Doc devoted his humble life to the care
and welfare of Asian elephants.
He enriched his colleagues by sharing
his peaceful wisdom and his penetrating
understanding of elephants.
His unwavering ethical convictions
are a legacy of hope for those
who carry the torch.
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CONTENTS
Foreword, by John Seidensticker / xi
Preface / xv
1. INTRODUCTION / 1
NEVER FORGETTING THE IMPORTANCE OF ETHICAL
TREATMENT OF ELEPHANTS
Christen Wemmer and Catherine A. Christen
PART I. OVERVIEW OF ELEPHANT PHILOSOPHY AND SCIENCE
2. ELEPHANTS IN TIME AND SPACE / 00
EVOLUTION AND ECOLOGY
Raman Sukumar
3. PERSONHOOD, MEMORY, AND
ELEPHANT MANAGEMENT / 00
Gary Varner
4. ELEPHANT SOCIALITY AND COMPLEXITY / 00
THE SCIENTIFIC EVIDENCE
Joyce H. Poole and Cynthia J. Moss
5. ELEPHANTS, ETHICS, AND HISTORY / 00
Nigel Rothfels
6. PAIN, STRESS, AND SUFFERING IN ELEPHANTS / 00
WHAT IS THE EVIDENCE AND HOW CAN WE MEASURE IT?
Janine L. Brown, Nadja Wielebnowski, and Jacob V. Cheeran
PART II. ELEPHANTS IN THE SERVICE OF PEOPLE: CULTURAL
DIFFERENCES AND ETHICAL RELATIVITY
7. ELEPHANTS AND PEOPLE IN INDIA / 00
HISTORICAL PATTERNS OF CAPTURE AND MANAGEMENT
Dhriti K. Lahiri Choudhury
8. CARROTS AND STICKS, PEOPLE AND ELEPHANTS / 00
RANK, DOMINATION, AND TRAINING
John Lehnhardt and Marie Galloway
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9. CANVAS TO CONCRETE / 00
ELEPHANTS AND THE CIRCUS-ZOO RELATIONSHIP
Michael D. Kreger
10. WHY CIRCUSES ARE UNSUITED TO ELEPHANTS / 00
Lori Alward
11. VIEW FROM THE BIG TOP / 00
WHY ELEPHANTS BELONG IN NORTH AMERICAN CIRCUSES
Dennis Schmitt
12. THE CHALLENGES OF MEETING THE NEEDS
OF CAPTIVE ELEPHANTS / 00
Jane Garrison
13. MOST ZOOS DO NOT DESERVE ELEPHANTS / 00
David Hancocks
14. ZOOS AS RESPONSIBLE STEWARDS OF ELEPHANTS / 00
Michael Hutchins, Brandie Smith, and Mike Keele
15. CAN WE ASSESS THE NEEDS OF ELEPHANTS IN ZOOS?
CAN WE MEET THE NEEDS OF ELEPHANTS IN ZOOS? / 00
Jill D. Mellen, Joseph C. E. Barber, and Gary W. Miller
16. GIANTS IN CHAINS / 00
HISTORY, BIOLOGY, AND PRESERVATION
OF ASIAN ELEPHANTS IN CAPTIVITY
Fred Kurt, Khyne U Mar, and Marion E. Garaï
PART III. ELEPHANTS AND PEOPLE IN NATURE: THE ETHICS OF
CONFLICTS AND ACCOMMODATIONS
17. RESTORING INTERDEPENDENCE BETWEEN
PEOPLE AND ELEPHANTS / 00
A SRI LANKAN CASE STUDY
Lalith Seneviratne and Greg D. Rossel
18. SUMATRAN ELEPHANTS IN CRISIS / 00
TIME FOR CHANGE
Susan K. Mikota, Hank Hammatt, and Yudha Fahrimal
19. HUMAN-ELEPHANT CONFLICTS IN AFRICA / 00
WHO HAS THE RIGHT OF WAY?
Winnie Kiiru
20. PLAYING ELEPHANT GOD / 00
ETHICS OF MANAGING WILD AFRICAN ELEPHANT POPULATIONS
Ian Whyte and Richard Fayrer-Hosken
viii CONTENTS
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21. TOWARD AN ETHIC OF INTIMACY / 00
TOURING AND TROPHY HUNTING FOR ELEPHANTS IN AFRICA
Rebecca Hardin
22. THE ETHICS OF GLOBAL ENFORCEMENT / 00
ZIMBABWE AND THE POLITICS OF THE IVORY TRADE
Rosaleen Duffy
Contributors
Index
ix CONTENTS
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4 ELEPHANT SOCIALITY
AND COMPLEXITY
THE SCIENTIFIC EVIDENCE
JOYCE H. POOLE AND CYNTHIA J. MOSS
Conventional wisdom places elephants among the more intelligent, socially
intricate, and emotionally complex nonhuman species. Hancocks (Chap-
ter 13 in this volume), citing Morris and Morris (1966) and Kellert (1989),
notes that people rank elephants among the “most-liked” animals and con-
sider the intelligence of elephants to be one of the most preferential fac-
tors. This popular conception reflects a long-term legacy of both legend and
research about elephants. Scientists and philosophers, going back to Aris-
totle, have claimed elephants to be highly intelligent, and some have even
viewed them as quasi-moral agents (Meredith 2001). This chapter exam-
ines what is currently known about the intelligence and complexity of ele-
phants and attempts to answer whether they are deserving of special moral
consideration.
Although the social complexity of elephants has been studied in depth,
little systematic research on their cognitive abilities has been carried out
until very recently (see Rensch 1956, 1957; Hart et al. 2001; McComb et al.
2001; Nissani 2004; Douglas-Hamilton et al. 2006; McComb, Bates, and
Moss 2006; Plotnik, de Waal, and Reiss 2006). Though this body of research
is growing rapidly, much of what is currently claimed regarding the supe-
rior intelligence of elephants is based on anecdotal evidence. Anecdotal ev-
idence falls into several categories, and as long as the nature of this evidence
is understood, some of it may be used to substantiate certain claims. For
example, much critical data related to elephant cognition are based on rare
behavior, which scientists have observed on an ad libitum (discretionary)
basis. The observations of these scientists, who are trained in the interpre-
tation of behavior, are an important data set (Byrne 1997a).
In this chapter, we provide empirical verification of elephant intelligence
and social complexity. We examine such factors as social structure and flex-
ibility, social network size, social learning, behavioral innovation, relative
brain size and complexity, memory, and communication. We discuss Machi-
avellian intelligence, insight, and theory of mind, mirror self-recognition,
and reaction to death. We argue that current and emerging evidence indi-
cates we should err on the side of caution and treat elephants with special
consideration. Evidence for these arguments comes in the form of scien-
tific publications, previously unpublished data, and scientific records of
69
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rare behaviors. To reduce speculation as to the validity of the accounts of
rare behavior, we rely on our own observations or those of our colleagues.
The previously unpublished data presented in this essay come primarily
from the Amboseli Elephant Research Project (AERP), a thirty-four-year
study of known individuals in Amboseli National Park, Kenya.
Three recognized species of elephants exist: Asian elephants, Elephas
maximus, African savannah elephants, Loxodonta africana, and African for-
est elephants, L. cyclotis. This chapter applies generally to all elephants. Al-
though conclusions regarding the behavior of wild elephants come pri-
marily from observations of African savannah elephants in Amboseli, and
conclusions from experimental studies of captive elephants are primarily
based on Asian elephants, it is generally acknowledged that there is broad
similarity among the three species (Payne 2003; Sukumar 2003). Except
where particularly relevant we have not attempted to compare elephants
with other species.
The point of this chapter is not to argue that elephants deserve un-
precedented rights that other species do not. Perhaps other animals also de-
serve such consideration, but we are not qualified to judge that. Many
mammals may equal or even surpass elephants in their individual abilities,
but we argue that it is the totality of elephant characteristics that makes
them deserving of special consideration.
This chapter relies on a common understanding of certain basic con-
cepts of elephant social structure. We define these here.
Family unit or family: One or more adult females and calves with a high
frequency of association over time, who act in a coordinated manner and
exhibit affiliative behavior toward one another (Moss and Poole 1983). This
term does not exclude two or more adult females without offspring, or a
single adult female and one or more juveniles who are not her immediate
offspring, making up a family. The term family and family unit may be used
interchangeably. In Amboseli, each family is referred to by a two-letter
code, such as AA, AB, and so on.
Bond group: Two or more family units who associate with one another
at high frequency relative to their associations with other family units in the
population and whose members display affiliative behavior toward one an-
other (Moss and Poole 1983).
Group: Any number of elephants of any age or sex moving together in
a coordinated manner with no single member or subgroup at a distance
from its nearest neighbor greater than the diameter of the main body of the
group at its widest point.
Aggregation: A group specifically made up of more than one family unit
with or without associating independent adult males.
Fission-fusion society: In relation to elephants the term fission-fusion so-
ciety refers both to the slow changes in the structure of families or bond
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groups that occur over the course of years or decades and the very rapid
changes that occur in social group composition over the course of hours.
In Amboseli, we have collected records on family group associations in
two basic ways: as sightings records and as census records. Sightings record
the presence of families, or portions of families, in any group that is en-
countered in the course of a day. Presence requires at least one member of
a family to be sighted. We maintain the records in a Microsoft Access data-
base (henceforth cited in this chapter as “AERP database”) that contained
more than 34,000 records as of the end of 2005. Censuses record the pres-
ence or absence of each family member at a sighting.
Social Complexity and Flexibility
Elephants live in a fluid, multitiered, fission-fusion society where group
membership changes frequently, forming and dividing along lines that may
be predicated on close social bonds, home range, and season (Douglas-
Hamilton 1972; Moss and Poole 1983; Sukumar 2003; Wittemyer, Doug-
las-Hamilton, and Getz 2005; Archie, Moss, and Alberts 2005; Moss and
Lee, forthcoming). Families are composed of a discrete, predictable com-
position of individuals, but over the course of hours or days, these groups
may temporarily separate and reunite or they may mingle with other so-
cial groups to form larger social units. The close and lasting social rela-
tionships formed by elephants are remarkable in the context of their fluid
social system (Archie et al. 2005; Moss and Lee, forthcoming). In fact, this
combination of social qualities—close and enduring cooperative social re-
lationships, and fission-fusion sociality—exists in only a small number of
cooperatively hunting carnivores (e.g., hyenas, lions, and sperm whales)
and also a few primates (e.g., chimpanzees and humans; Archie et al. 2005;
Moss and Lee, forthcoming).
Relationships radiate out from the mother-offspring bond through
members of the family, bond group, clan, subpopulation, to independent
adult males and even beyond the population to strangers. Although spe-
cial relationships between individual elephants may last a lifetime, the qual-
ity of relationships and the structure and degree of cohesion in an indi-
vidual’s social network may also change through time. With a maximum
lifespan in the wild of sixty-five years (Moss 2001; AERP database), ele-
phants are unusually long-lived mammals (Eisenberg 1981) and thus their
relationships are extremely long term.
Like many sexually dimorphic mammals, adult male and female ele-
phants live in very different social worlds (Moss 1988; Poole 1994). A com-
plex network of bonds between individuals and families characterizes the
lives of females and their offspring, while fluctuating sexual cycles distin-
guish the dynamic activities, associations, and relationships of males (Moss
and Poole 1983). Elephants show strong individual personalities that affect
71 ELEPHANT SOCIALITY AND COMPLEXITY
—forthcoming.
[FIGURE] Figure 4.1
about here
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how they interact with other elephants and how well they are able to in-
fluence members of their group (Moss and Lee, forthcoming).
Elephant Social Structure
Members of an elephant family exhibit a high frequency of association over
time, display strongly affiliative behavior, including a pattern of greeting cer-
emonies (Moss 1981) and are highly cooperative in group defense, resource
acquisition, offspring care, and decision making (Douglas-Hamilton 1972;
Dublin 1983; Moss and Poole 1983; Lee 1987; Moss 1988; Poole 1998; Payne
2003). A matriarch, usually the oldest female, leads each family, and most,
though not all family members, are genetically related. Archie et al. have
found complete uniformity of mitochondrial DNA (mtDNA) haplotypes in
approximately 90% of Amboseli families. Average pair-wise relatedness be-
tween adults in families is 0.14, similar to the value expected in first cousins
(r= 0.125) (Archie et al. 2005). Over thirty-four years of study in Amboseli,
through 2004, there have been sixty-one families, seven of which no longer
exist because of the deaths of their members. In the same period of time,
new family units have been formed by five fissions and two fusions. Family
membership has ranged from two to fifty-two individuals and, in 2002, av-
eraged 18.7 individuals (Moss and Lee, forthcoming).
72 OVERVIEW OF ELEPHANT PHILOSOPHY AND SCIENCE
Figure 4.1 Families may temporarily separate and reunite or they may mingle
with other social groups to form larger social units. The close and lasting social
relationships formed by elephants are remarkable in the context of their fluid
social system.
Photograph courtesy of Petter Granli / ElephantVoices.org
—forthcoming.
[FIGURE] Fig-
ures 4.2 and 4.3
about here
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73 ELEPHANT SOCIALITY AND COMPLEXITY
Figure 4.2 An African (top) and Asian (bottom) elephant family group walks
in close proximity, providing care and reassurance to infants and calves.
Photograph of African elephants courtesy of Petter Granli / ElephantVoices.org;
photograph of Asian elephants courtesy of Joyce Poole / ElephantVoices.org
1.Wemmer,Elephants and Ethics 2/1/08 2:23 PM Page 73

74 OVERVIEW OF ELEPHANT PHILOSOPHY AND SCIENCE
Figure 4.3 An Asian male elephant in musth tests females for receptivity (top);
two African elephant males in musth duel for supremacy and access to an
estrous female (bottom).
Photograph of Asian elephants courtesy of Petter Granli / ElephantVoices.org;
photograph of African elephants courtesy of Joyce Poole / ElephantVoices.org
1.Wemmer,Elephants and Ethics 2/1/08 2:23 PM Page 74

Above the level of the family unit a second tier of relationships exists
within bond groups (Douglas-Hamilton 1972; Moss and Poole 1983; Wit-
temyer et al. 2005). A bond group may include as many as five families
(Moss 1988). Although the ties between individuals at this level are weaker
than those within a family, bond group members also exhibit greeting cer-
emonies, form alliances against aggressors, assist in the care of one an-
other’s offspring, and defend one another in times of danger. Since the ma-
jority of bond groups are probably fission products of former families
(Moss 1988) most carry the same mtDNA haplotype (Archie et al. 2005).
Yet, rarely, there do exist bond groups (closely cooperating individuals)
composed of unrelated families. Archie et al. (2005) have pointed out that
even within families there are individuals who, through chance demo-
graphic events, have no close relatives. Yet, these individuals still benefit
from the same cooperative behavior. Cohesion within families and bond
groups varies significantly and depends upon a combination of factors, in-
cluding individual personalities, the formation and dissolution of individ-
ual social bonds, the strength of the matriarch’s leadership, historical
events such as deaths of influential individuals, the type of habitat, and the
season (Moss and Lee, forthcoming).
Clans are the next social level and have been defined as families who
share the same dry-season home range (Laws and Parker 1968; Douglas-
Hamilton 1972; Moss and Poole 1983). A clan is usually composed of
several bond groups and numerous families, such that several hundred
elephants may make up a clan. Playback experiments using individual
vocalizations (McComb, Moss, Sayialel, and Baker 2000) indicate that ele-
phants, too, distinguish between more familiar associates (typically clan
members) and less familiar ones (typically nonclan).
Many of Amboseli’s elephants are still in the same family, occupying
the same home range that they were thirty-four years ago, while some
families have shifted clan, or exchanged bond group. A few individuals
whose families have gone extinct have been adopted by unrelated fami-
lies, and in one case, individuals abandoned their natal family to create
a new family with members of a previously unrelated family (Moss 1988;
AERP database). Long-term records demonstrate both the extraordinary
fidelity and unusual flexibility of elephant social relationships (Moss and
Lee, forthcoming).
During dry conditions, families tend to move in relatively small groups
in their clan area, but when resources are abundant, families may gather in
large aggregations in an expanded range (Poole and Moss 1989). In these
aggregations, several clans may intermingle. Far from being random, the
fission and fusion of elephant groups follow predictable patterns, tending
toward association according to hierarchical clustering into families, bond
groups and clans, and breaking down in the reverse order. Elephants ag-
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gregate for different purposes (Western and Lindsay 1984), including an-
tipredation (Laws, Parker, and Johnstone 1975), social benefits (Moss
1981), and improved mating opportunities (Poole and Moss 1989).
Although many mammals may follow some of the patterns exhibited by
elephants, the combination of enormous social fluidity and durability of
elephant associations and relationships is remarkable and rivals that of
chimpanzees and humans in its complexity (McComb et al. 2000; Archie
et al. 2005; Moss and Lee, forthcoming). Also notable is that kinship is not
the only determinant of social bonds. Female elephants with few kin are not
excluded from the benefits of sociality.
Young male elephants grow up in the tightly bonded society of females,
maintaining close relationships with their relatives and participating in the
many social events that affect their family, albeit at a lower intensity than
their female agemates (Lee and Moss 1999; Poole, forthcoming). Males de-
part from their natal families between 9 and 18 years of age (Lee and Moss
1999). A newly independent male must acquire a fresh set of behaviors to
adapt to the society of males, where body size and fluctuating sexual state
determine interactions and relationships (Poole 1989a). His transition
from one society to the other occurs gradually, but dramatically, over a
period of several years (Lee, Poole, and Moss, forthcoming). During this
time, a young male spends much of his time getting to know his agemates,
and sparring and playing with novel partners from outside his natal fam-
ily (Lee 1986). Thus he gathers information crucial to his longevity and re-
productive success (Poole 1989a, 1989b; Lee et al., forthcoming; Poole, Lee,
and Moss, forthcoming). Once fully independent, he forms relatively few
close and long-lasting bonds with other elephants. Though males are often
seen in small, all-male groups and may form lasting associations with cer-
tain individuals, these are rather loose arrangements (Croze 1972; Lee et al.,
forthcoming). During sexually active periods males rove from one family
group to the next, using olfactory and acoustic cues to search for receptive
females (Poole and Moss 1989).
Elephants Have an Unusually Large Social Network
The social complexity hypothesis proposes that high intelligence has
evolved to help individuals anticipate the plastic and often unpredictable
behavior of group members (Jolly 1966). Social complexity has been cited
as one indicator of intelligence (Byrne and Whiten 1988), and group size
has been used as a measure of social complexity (Dunbar 1992). Group in
this sense refers to long-term associates whose characteristics and histories
are likely known to one another, rather than a temporary aggregation of
strangers. This concept works well for species that live in stable, imperme-
able groups, but less well for species like humans, chimpanzees, whales, and
elephants, that do not. For fission-fusion societies the term “social net-
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work” encompasses such long-term associates. Just how large is an ele-
phant’s social network?
The Amboseli population is relatively small (1,417 at December 2005)
compared with other elephant populations; yet, it is, nonetheless, a large so-
ciety (McComb et al. 2000). In Amboseli, a female elephant may encounter
literally hundreds of other individuals in the course of her daily range, and
34 years of records show that each family has been found with every other
family in the population at least once (Moss and Lee, forthcoming).
Taking Amboseli’s GB family as an example, we may estimate the size
of one female’s social network. Between 1974 and 2002, the GB family has
grown from fourteen to thirty-seven members. Over this time, any given
member of Amboseli’s GB family found herself in a median group size of
thirty-nine individuals, although at any particular moment she might
(rarely) have been alone, with a small portion of her family, or in a group
of 550 individuals (interquartile range 25–95; N= 1,294; AERP database).
Members of the GB family have been sighted without other families on 428
occasions and with other families (in aggregations) 866 times. Of these 866
sightings, 59% of the aggregations included the IB family. The GB family
was seen in aggregations with fifty other families, too, of which fourteen
particular families were present in 15–30% of the sightings. Taking the pop-
ulation’s current average family size, these close associates account for an
estimated 261 elephants of which 125 (48%) are adult females at least ten
years of age.
The question remains, however, do these animals know one another as
individuals? To show that these are not just temporary aggregations of
strangers, we must prove that elephants are able to distinguish between in-
dividuals, and to retain this knowledge over time. Work by biochemist Ras-
mussen and veterinarian Krishnamurthy (2000) found that offspring
flehmen (a “chemical analysis” accomplished by touching the tip of the
trunk on a liquid substance and placing it against the vomeronasal organ
on the roof of the mouth) twice as often when presented with their
mother’s urine than with any other individual’s urine, suggesting that they
are able to distinguish their mother’s urine. Even after a twenty-seven-year
separation an offspring showed a flehmen response to its mother’s urine
(Rasmussen and Krishnamurthy 2000). Elephants answer the contact calls
of close relatives, and, working in Amboseli, ethologist McComb et al.
(2000) found that playing back the contact call of an elephant who had died
elicited contact calling by her family unit almost a year after her death. Mc-
Comb et al. (2000) also found that elephants recognized and usually an-
swered the individual voices of their own family and bond group (ca. 40–
60 individuals), and they also distinguished between familiar and less
familiar elephants on the basis of how often they were encountered. Mc-
Comb et al. (2000) predicted that the females would have to be familiar
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with the voices of at least 100 adult females to make the observed discrim-
inations, indicating that elephants have one of the largest known social net-
works of any nonhuman species.
Males, too, make use of social knowledge in a large network. Comple-
mentary male postures in long-distance dyadic interactions (i.e., those be-
tween a paired group of individuals) indicate that males recognize one an-
other visually, and that they are aware of one another’s relative size, strength,
and condition (Poole 1982, 1989a). An instance of such interaction would
be a higher-ranking male posturing with Head-High; Ear-Spreading; Ear-
Folding; Advance-Toward, while the corresponding lower-ranking male is
posturing with Head-Low, Ear-Flattening; Turn-Away; Retreat-From; (Poole
1987, 1999a; Kahl and Armstrong 2000; Poole and Granli 2003). Males also
use sound (Poole et al. 1988; Poole and Moss 1989; Langbauer et al. 1991)
and scent (Rasmussen et al. 1982; Rasmussen and Schulte 1998), to remotely
monitor the location of receptive females. Approximately, three weeks be-
fore true estrus and ovulation, female elephants exhibit a period of estrous-
like behavior, referred to as “pre-estrus” (Poole and Moss 1989), that is as-
sociated with an anovulatory LH (lutenizing hormone) surge (Kapustin et
al. 1996) and increasing pheromone secretions (Rasmussen, Krishnamurthy,
and Sukumar 2005). Males accompanying a female during the pre-estrus are
likely to be the same individuals who associate with her later during her true
estrus (AERP database). During the intervening days, these same males may
be many kilometers apart from that female, searching for, or consorting
with, other females (AERP database). The median number of males in as-
sociation with an estrous female in Amboseli is 5 (interquartile range 3–8;
range 1–40; N= 728; AERP database). Because upward of 100 adult males
may associate with female groups in the population at any given time, there
is strong suggestion that individual males may relocate a female during true
estrus using acoustic and olfactory cues as well as memory of her individ-
ual identity and clan area, and therefore of her likely location.
Although other mammals are also able to recognize the voices and scents
of conspecifics (members of their own species), the sheer number and long-
term retention of social information makes elephants unusual. McComb
et al. (2001) showed that such vocal discriminations are learned through
years of experience and older individuals are more discerning than younger
ones. McComb et al. (2000) argue that an unusually large and fluid social
network may be a phenomenon unique to long-lived mammals like ele-
phants, equipped with long-range signaling capability and the mental ca-
pacity for extensive social recognition.
Rate of Change in Social Setting
Given a large and fluid social network in which group composition changes
rapidly, elephants may encounter literally hundreds of individuals each day.
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For example, of the 1,294 sightings of the GB family, 311 (24.0%) were of
groups of 100 individuals or more; 190 (14.7%) were of groups of 150 ele-
phants or more, and 122 (4%) were of groups of 200 elephants or greater.
The number of elephants encountered by each female each day, and the
rate of change, prove unusually large, especially when one considers that
these data represent instantaneous sightings and that groups change in size
and composition many times per day (Mutinda 2003).
Sexually active males, searching from group to group for receptive fe-
males, may also meet and interact with hundreds of different individuals,
male, and female, in the course of a day (Poole and Moss 1989). The na-
ture of a male’s interactions with those he meets is strongly influenced by
his age, body condition, and sexual state (Poole 1982, 1989a).
Elephants are able to recognize the voices, scent, and appearance of
many individuals, some of whom they encounter only relatively rarely. Al-
though the social setting of many other species may also change during the
course of a day, the sheer number of individuals involved and the hierar-
chical nature of the formation and dissolution of aggregations make ele-
phants unusual. Under these circumstances, the importance of being able
to distinguish genuine strangers from a wide range of more regular asso-
ciates is not insignificant (McComb et al. 2001).
Social Learning
There is empirical evidence that social learning and behavioral innovation
are positively correlated with brain size in mammals (Reader and Laland
2002). Both behavioral innovation and social learning are essential ele-
ments in the development and maintenance of elephant social complexity.
Learning What to Eat
Infant elephants must learn and practice complex coordination of the
trunk and apply this gradually acquired skill to the procurement of ap-
propriate foods (Lee and Moss 1999). Calves begin to sample potential food
items when they are 1 to 2 months old. Sampling consists of grasping ob-
jects with their trunk, rolling food items in their trunk, and placing food
matter in their mouths. At times young calves may chew, but sampling
starts well before actual ingestion. Calves gradually acquire foraging knowl-
edge by sampling what adults are eating. They attain this information by
placing their trunks in the mouths of adults and pulling out food items,
stealing food from their mothers and allomothers, as well as by eating the
fresh dung of other elephants (Eisenberg 1981). Throughout their first five
years, calves spend 15.8% of their social contact time exploring the food in-
take of others (Lee and Moss 1999). Elephants have many food sources, and
social learning allows calves to exploit a wide range of these seasonally and
geographically varying species (Lee and Moss 1999).
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Learning to Care for Calves
Juvenile females play a vital role in the care of young calves (Lee 1987). Allo -
mothering both increases calf survival (Lee and Moss 1986) and provides
young females with an array of caretaking experiences that persist until they
give birth to their own first calf). Despite this practical social experience,
80 OVERVIEW OF ELEPHANT PHILOSOPHY AND SCIENCE
Figure 4.4 Allomothering increases calf survival and provides young females
with an array of caretaking experiences that persist until they give birth to
their own first calf. A juvenile female rescues her younger sibling from an at-
tempted kidnapping by a nonfamily juvenile female and calls for the assis -
tance of her mother (top); juvenile females shepherd infant Emily Kate back
to her mother (bottom).
Photograph courtesy of Petter Granli / ElephantVoices.org
1.Wemmer,Elephants and Ethics 2/1/08 2:23 PM Page 80

first-born infants still have higher mortality rates than infants born to ex-
perienced mothers (Moss 1994), indicating that even by the age of first par-
turition (14–15 years) female elephants still have much to learn about
mothering. The calves of inexperienced mothers show higher levels of dis-
tress than do calves born to experienced mothers, who appear to be more
responsive to calf demands for food and protection, with obvious conse-
quences for calf growth and survival (Lee and Moss 1999). In addition,
first-time, or primiparous, mothers are given more assistance from family
members than are older mothers. Knowledge gained from experiences over
a succession of births plays an important role in calf survival.
Social Learning as an Aspect of Female Reproductive Behavior
Some behaviors essential to mate choice, such as consort behavior, appear
to require a social context for learning. Young estrous females do not
demonstrate the estrous behavior typical of older females and are fre-
quently chased and mounted by a succession of young and nonmusth
males (Moss 1983). As a female gains individual experience through suc-
cessive estrous periods her responses to males change; she learns to avoid
young males and to select and go into consort only with large musth males
(Moss 1983; Poole 1989b). Both the acquisition of estrous behaviors and
the choice of mates appear to be facilitated by the presence and behavior
of the mothers of these young females (AERP database). Mothers of young
estrous females are often observed exhibiting estrous postures and behav-
iors when not in estrus themselves (AERP database; J. Poole, personal ob-
servation). Mothers (and occasionally nonmother matriarchs) may be ob-
served to approach and avoid males, run with their young estrous daughters
(or young estrous members of their family) during long chases and occa-
sionally make postcopulatory calls after the young female is mated (BBC
Natural History Unit 1995). The behavior of mothers and daughters dur-
ing a daughter’s first estrous period not only indicates the importance of a
social context for learning, but also suggests that mother elephants may be
engaged in a rudimentary form of true teaching.
The birth of a female’s first calf is another documented event where the
presence and behavior of experienced females aids inexperienced mothers.
Experienced family members assist young females to cope with the physi-
cal demands of birth, including helping a newborn to its feet, and with the
immediate protection and socialization of the newborn calf (Moss 1988;
Lee and Moss 1999).
Social Learning as an Element in Male Independence
In exploratory and play behavior, young males seek same-sex play partners
outside the family, thereby socializing and bonding with their future peer
group while simultaneously assessing the strength of their future rivals (Lee
81 ELEPHANT SOCIALITY AND COMPLEXITY
—survival.
[FIGURE] Figure
4.4 about here
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1986). This male behavior can be linked to male emigration from the na-
tal family, an event that occurs at an average age of 14 years (Lee and Moss
1999). The process of a male’s departure from his family averages 16
months, but ranges anywhere from 5 to 35 months (Lee et al., forthcom-
ing). The wide range in duration indicates that the process of male inde-
pendence is probably a highly individualized event. Male mortality in the
10–20-year age bracket is also significantly higher than in females (Moss
2001). To survive on their own newly independent males require sufficient
social experience with other males along with knowledge of male areas of
residence and surrounding habitat (Lee and Moss 1999).
Social Learning and Role Models in Male Reproductive Behavior
Young males appear to learn to distinguish between reproductive odors. For
example, young males, though not older males, often mistake the smell of
a pre-estrous female or one who has just given birth, for that of a receptive
female (J. Poole, personal observation). Successful mounting and intro-
mission, too, appears to require considerable skill and experience. This
learning process may, in part, be gained by watching the behavior of older,
more experienced males. Young male elephants are often observed to fol-
low older musth males, testing the same urine spots and females that they
do. Musth males are extremely tolerant of these youngsters, allowing them
to stand less than a meter from an estrous female while older males are kept
at long distances (Poole 1982). Experience from southern Africa also high-
lights the importance of social learning in the acquisition of appropriate
male reproductive behavior (Slotow et al. 2000). Juvenile male elephants
that experienced their families killed in a culling operation and were then
introduced to areas without adult role models exhibited abnormal repro-
ductive behavior as young adults, including the mounting, tusking and
killing of black rhinos. While trauma is likely to have been a causal factor
in the development of this abnormal behavior (Bradshaw et al. 2005), it is
also likely that the absence of adult males contributed to the inappropri-
ate sexual reaction of these young males.
The Role of Social Learning in Tool Manufacture and Use
Elephants are unusual among nonprimates in that they use and even man-
ufacture simple tools. The sensorimotor specializations of the trunk are ex-
tensive (Rasmussen and Munger 1996), allowing delicate manipulations of
both large and small objects. Elephants pick up objects (logs, rocks) and
throw them at opponents, use sticks to remove parasites, branches as fly
switches, and logs to neutralize electric fences (Poole 1998). A study of
Asian elephants by ethologist and sociologist Lynette Hart and colleagues
(2001) indicates that elephants modify the length of long branches to make
a suitable fly switch. That study also describes the role of social learning in
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the acquisition of tool use. Adult Asian elephants modif y branches by
pulling off leaves and shortening sticks to create fly switches of ideal
length. Two young elephants, ages 18 and 9 months, imitated the fly
switching behavior of adults. The 18–month-old calf was able to remove
a side branch successfully and was coordinated in switching with the mod-
ified shorter branch. The 9–month-old showed only uncoordinated at-
tempts to switch, but her movements appeared to imitate the fly switching
behavior of older elephants.
Vocal Learning
Very few mammals are capable of producing or modifying sounds in re-
sponse to auditory experience (McCowen and Reiss 1997; Tyack and
Sayigh 1997; Janik and Slater 2000). Those that can include humans, some
marine mammals (e.g., bottlenose dolphins, Tursiops truncates; harbor
seals, Phoca vitulina; humpback whales, Megaptera novaeangliae) and bats.
Despite the obvious significance of vocal imitation for human-language ac-
quisition, there is scant evidence for vocal production learning in other pri-
mates. Recently, Poole et al. 2005 provided two examples of vocal imita-
tion in African elephants: A young female elephant who imitated the
sounds of trucks, and an African male zoo elephant who imitated the chirp-
ing sounds of the Asian elephants with whom he was raised. Earlier reports
have described Asian elephants who have learned to produce whistling
sounds by blowing air through their trunks, and still other elephants who
have imitated this innovative whistling technique (Wemmer and Mishra
1982; Wemmer, Mishra, and Dinerstein 1985; I. Douglas-Hamilton, per-
sonal communication). Vocal production learning enables a flexible and
open communication system, involving both behavioral innovation and
social learning, in which animals may learn to imitate sounds not typical
of their species. Poole et al. 2005 suggest that vocal learning should likely
occur in other species in which long-lived social bonds are based on indi-
vidual-specific relationships and involve fluid group membership, and
where vocal communication functions in maintaining contact, in individ-
ual or group recognition and in mediating social interactions.
One of the hypotheses of the origin of human language is that it arose to
encode increasingly complex information about social relationships (Wor-
den 1998; Dunbar 2003; Pinker 2003). The association between species that
show vocal production learning abilities and those with fission-fusion soci-
eties supports this hypothesis (e.g., cetaceans, elephants, chimpanzees, hu-
mans). In describing the social origin of language hypothesis, Pinker (2003),
has suggested that sociality, knowledge gathering, and language co-evolved
in humans, in that language reinforces social relationships and provides a
mechanism for distributing knowledge to associates. These three character-
istics of human societies appear to be evident, to a lesser extent, in elephants.
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Communication as a Measure of Social and Cognitive Flexibility
Species that make use of both contextual learning (behavioral context and
sequential positioning) and production learning (modification based on ex-
perience) are theoretically capable of developing a more complex acoustic
communication system than species utilizing only the former. Each addi-
tional form of vocal learning can increase the complexity of a communi-
cation system; the more forms present the greater the system’s potential for
openness and plasticity (Janik and Slater 2000).
Elephants make use of both contextual and production learning and,
as predicted, their acoustic communication system includes an extensive
vocal repertoire with a high degree of variability both within and between
individuals (Poole et al. 1988; Langbauer 2000; McComb et al. 2000; Leong
et al. 2003; Soltis, Leong, and Savage 2005a, 2005b; Poole, forthcoming).
Elephants also have a wide range of visual and tactile gestures and displays
(Kahl and Armstrong 2000; Poole and Granli 2003) and intricate chemi-
cal communication (Rasmussen and Krishnamurthy 2000). Together this
complex suite of vocal, chemical, visual, and tactile signals mediates the in-
tricate teamwork displayed by members of an elephant family (Poole,
84 OVERVIEW OF ELEPHANT PHILOSOPHY AND SCIENCE
Figure 4.5 Members of an elephant family exhibit a high frequency of associa-
tion over time and display strongly affiliative behavior, including a pattern of
greeting ceremonies. Here mother and daughter greet one another.
Photograph courtesy of Petter Granli / ElephantVoices.org
1.Wemmer,Elephants and Ethics 2/1/08 2:23 PM Page 84

forthcoming; Poole and Granli 2003). Day to day decision making (e.g., de-
ciding when and where to go; antipredator responses) involves broad vo-
cal participation and often includes the building of consensus through vo-
cal exchanges that may take up to an hour (J. Poole, personal observation;
C. Moss, personal observation).
Indicators of Elephant Intelligence
Brain Size and Complexity
Until recently, only scant information was available on the brains of ele-
phants, with the majority of the literature based on only a few specimens
(Cozzi, Spagnoli, and Bruno 2001). Recent work has added to our knowl-
edge (Hakeem et al. 2005; Shoshani, Kupsky, and Marchant 2006). The
brains of Asian and African elephants rank among the highest of all ani-
mals for absolute and relative mass and cortical expansion and complex-
ity, features comparable only to those of some of the cetaceans, the great
apes, and humans. Weighing 4.5–6.5 kilograms, the elephant’s brain is the
largest in absolute mass among land mammals (Cozzi et al. 2001), with the
brain of E. maximus weighing up to 5.5 kilograms, and that of L. africana
being slightly heavier and larger. The temporal lobes of the elephant’s brain,
which are thought to function in recognition, storage and retrieval of in-
formation related to sight, touch, smell, and hearing, are especially large
and extremely complex (Shoshani 1998; Shoshani et al. 2006).
The encephalization quotient (EQ), the ratio between the observed and
expected brain weight for a defined body weight (Jerison 1973), implicitly
holds that body size is taken as a “given” and selection operates only on
brain size. An EQ equal to 1.0 is an “average” mammalian brain. Primates
all have relatively high EQs, with Homo sapiens at 7.0+ demonstrating a
quotient far above that of all other mammals. Elephant values, at between
1.7 (Shoshani 1998; Eisenberg 1981) and 2.3 (Cutler 1979) are compara-
ble to those of larger primates (e.g., chimpanzee: 2.2–2.4; gorilla: 1.4–1.7;
orangutan: 1.6–1.9; Eisenberg 1981). Where sexual dimorphism is pro-
nounced, as in elephants, males have lower EQ values than females (EQ L.
africana: male: ca. 1.0; female: 2.0; Eisenberg 1981). Clearly age and sex are
important criteria in determining the importance of EQ, and for a variety
of reasons many have argued that EQ may not be a particularly useful guide
to relative intelligence (e.g., Byrne 1996; Roth 1999). Neurobiologist Roth
(1999) and cognitive ethologist Byrne (1996) argue that what should be im-
portant for intelligence is the absolute number of nerve cells in the brain,
as more nerve cells mean the potential for more complicated networks. Ele-
phants have a very thick cerebral cortex, and although its cell density is
lower than that of humans, elephants are estimated to have as many neu-
rons as humans, namely between 1011 and 1012. Shoshani et al. (2006) pos-
85 ELEPHANT SOCIALITY AND COMPLEXITY
—observation.
[FIGURE] Figure
4.5 about here
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tulate that convergent evolution as seen in complex learned skills and be-
havior may be responsible for the many similarities observed between the
human and elephant brain.
Long-Term Memory
As a general rule vertebrate species with absolutely large brains have de-
veloped the neocortex (the complicated parts of the cerebral cortex) to a
greater degree, have greater capacity for learning, and seem to be able to
learn more complicated tasks than vertebrates with absolutely smaller
brains (Rensch 1956, 1957). In addition, it seems that larger animals with
large brains also have the ability to retain information for longer than
smaller animals with smaller brains. In other words they have better mem-
ory (Rensch 1956, 1957). Exceptionally large and long-lived, elephants ac-
cumulate and retain social and ecological knowledge, remembering other
individuals (McComb et al. 2000; Rasmussen and Krishnamurthy 2000;
McComb et al. 2001) and places (Viljoen 1990; Shoshani and Eisenberg
1992) for years.
In experimental trials, elephants show an excellent ability to learn and
remember a large set of visual symbols and acoustic tones or commands
over long periods (Rensch 1957). Playback experiments in Amboseli also
provide good evidence for an elephant’s exceptional memory. Companions
use powerful calls to stay in contact when visually separated (Poole et al.
1988). As mentioned previously, elephants remember the contact calls of
family and bond group members and distinguish them from those of fe-
males outside these categories; moreover they can also discriminate be-
tween the calls of family units farther removed than bond group members
on the basis of how frequently they have encountered them (McComb et
al. 2000). In fact, McComb et al. (2000) predicted that females would have
to be familiar with (i.e., remember) the contact calls from at least 100 adult
females in order to perform these discriminations. McComb et al. 2001 also
found that older matriarchs possessed enhanced socially relevant, dis-
criminatory abilities, which are learned and remembered over many years
of experience. These older females acted as a repository of social knowledge
that contributed to increased fitness for the entire family.
In addition, playback to her family unit of the contact call of a 15–year-
old female that had died elicited contact calling three months after her
death, and contact calling and approach to the loudspeaker twenty-three
months after her death (McComb et al. 2000). Also, playback of the con-
tact call of a female who had departed from her family to join another one,
twelve years previously, also elicited contact calling (McComb et al. 2000).
Accounts of rare events by qualified observers also indicate that ele-
phants are able to remember the voice and perhaps scent of individual
people for over twelve years (R. Moore, personal communication; J. Poole,
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personal observation) and individual elephants for over 23 years (C. Buck-
ley, personal communication). In the latter case, the specific protective be-
havior of elephant Shirley (ca. 53 years) toward Jenny (ca. 30 years), when
reunited, suggested that they not only remembered one another, but also
remembered the adult-juvenile relationship they had once shared. In the
mid 1980s, one of us (J. Poole) established a relationship with Vladimir, a
wild male elephant, who sometimes came to her car window allowing her
to touch him (Poole 1996). In 2003, after a twelve-year separation, Poole
met Vladimir, then aged 34 years, and called to him. He walked up to her
window and allowed her to touch him. To the best of our knowledge, he
has not interacted with any other person in this manner (AERP colleagues,
personal communication).
Machiavellian Intelligence In Elephants
The term Machiavellian intelligence was coined to refer to the hypothesis
that increased intelligence was linked to the evolution, through competitive
tactics, of socially complex behavior predicated upon a good memory for
socially relevant information (Byrne and Whiten 1988; Byrne 1996). The
principles of Machiavellian intelligence were originally used to explain the
socially complex behavior of monkeys and apes, but more recently, non-
primate species have been shown to exhibit similar traits (see Byrne 1996).
Typical social traits include the formation of long-term social relationships,
use of third party or triadic intervention to decide the outcome of compet-
itive encounters, dependence on a network of allies, grooming used to build
up a network of support, active reconciliation used to repair strained rela-
tionships, use of tactical deception, and the classification of others by dom-
inance rank, affiliation or group membership. Successful use of these social
tactics demands good perception and discrimination, attention to social at-
tributes and good memory, all of which are displayed by elephants. The abil-
ities ascribed to Machiavellian intelligence require neocortical enlargement
and exceptional memory to retain and process socially relevant information
(Byrne 1997b), qualities that are, again, characteristic of elephants.
Elephants are contenders for rank among those species possessing
Machiavellian intelligence, in every respect. The formation of long-term re-
lationships is the very essence of elephant society and a network of allies,
who are usually though not always genetically related, defines the elephant
family and bond group. These supportive relationships are maintained not
by social grooming as in primates (Dunbar 1988), but through an elabo-
rate system of vocal, visual, chemical and tactile signals and ceremonies be-
tween relatives and friends (Moss 1988; Poole et al. 1988; see also Poole and
Granli 2003, 2004). The most well known of these is the greeting ceremony
(Moss 1988), but reunions are by no means the only situation under which
such a display of “bonding” occurs. Similar exuberant demonstrations of
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the importance of a social bond occur in a multitude of different contexts,
including, among others: after a family member’s infant or calf has been
lost, frightened, or denied access to the breast; if a member of the family
has been chased or harassed by another elephant; when a baby is born; or
following a disturbing or exciting event (Poole, forthcoming).
When key relationships are put at risk by conflict, some primates make
efforts to reconcile, and dyads that reconcile are more likely to continue to
support one another (Cords 1997; de Waal and van Roosmalen 1979). Sim-
ilarly, conflict between two individuals in an elephant family typically results
in vocal reconciliation that—even more sociably—usually involves the in-
tervention of a third party (J. Poole, personal observation). Third parties play
an important role in deciding the outcome of conflicts among elephants,
including support given to genetically unrelated individuals (Moss 1988;
Payne 2003 Archie et al. 2005). These “friendships” are maintained by re-
ciprocal altruism (Payne 2003). Work by McComb et al. (2000) shows clearly
that elephants are able to recognize and respond appropriately to the voices
of others and that they are able to classif y others by group membership at
the very least. Finally, there are many accounts of tactical deception by ele-
phants, though these are mostly anecdotal reports by elephant keepers
rather than data based on experimental trials or observations in the wild (e.g.,
see Rensch 1957; Chadwick 1992; Poole 1998). Many of these extraordinary
accounts are of elephants deceiving their keepers to obtain food or freedom.
Detailed documentation of such tactical deception is now needed.
Anticipatory Planning, Insight, and Theory of Mind
The combination of longevity, long-term memory, social learning, and be-
havioral innovation might lead us to expect elephants to display other cog-
nitive abilities, such as insight and theory of mind. Insight refers to be-
havior that shows the understanding of relations between stimuli and
events, while theory of mind is defined as an ability to understand that
others see, feel, and know (see also Varner, Chapter 3 in this volume).
In the wild, anecdotal evidence certainly suggests the use of insight by
elephants in response to electric fences. Elephants frequently disable elec-
tric fences by dropping logs onto them, or by uprooting and pushing trees
onto them, causing live wires to sag onto ground wires, thus shorting the
fence (Laikipia Ranch, Kenya, fence maintenance crew, personal commu-
nication; Seneviratne and Rossel, Chapter 17 in this volume). Reports exist
of elephants lifting their infants over fences (Chadwick 1992; Ngulia Sanc-
tuary, Tsavo National Park, Kenya, fence maintenance crew, personal
communication). In captivity, elephants have been known touse sticks to
pull out-of-reach food closer; open faucets, and, once these were bolted
shut, use rocks to break the nuts loose from the bolt; throw tires onto
nearby branches to weight them down to a level within reach; make a pile
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of tires to stand on so as to reach branches otherwise out of grasp (Chad-
wick 1992). Shoshani and Eisenberg (1992) recount an extraordinary tale
of an elephant that ingeniously placed vegetation under his feet to prevent
himself from sinking into muddy ground where he was tied, as he could
not reach dry ground. Evolutionary biologist Bernhard Rensch’s (1957) ob-
servations of the ability of domesticated elephants to work with minimal in-
struction and their talent to function as a team with extraordinary balance
and coordination, pushing and dragging heavy logs up inclined bars onto a
truck, caused him to credit elephants with true insight or “ideation”— the
ability to anticipate what will come of certain actions.
Behavior that qualifies as insight cannot be genetically wired or acquired
by trial and error. The problem must be new, or the animal must come up
with a new solution. The correct behavioral sequence must be arrived at
suddenly and completely and carried out relatively smoothly, with all its
constituent elements purposefully aimed at a single goal (Nissani 2004).
In Amboseli one of us (J. Poole) also witnessed behavior that meets
these criteria. In 1999 Poole witnessed Ella giving birth, an event accom-
panied by thirty minutes of excited vocalizations by her family and other
females (Poole 1999b). The commotion attracted other elephants, includ-
ing young and inexperienced Ramon, who upon sniffing Ella, mounted her,
his body and feet suspended above the newborn. Matriarch Echo, and her
adult daughter, Erin, rushed immediately to Ella’s side, backing, in paral-
lel, purposefully toward her and placing themselves resolutely on either side
of her. In what appeared to be a very deliberate attempt to prevent the male
from crushing the infant when he dismounted, their behavior appeared to
be a case of true insight. Both authors also witnessed Victoria and Virginia
(the two oldest females in a large family) rush to stand on either side of
Vega, who had been darted with an immobilizing drug, and push into her,
thus holding her up and preventing her from falling down. Only after force-
ful driving at them with vehicles would they give up.
Theory of mind accounts exist, too, with an often-related tale being that
of Chandrasekhan, the elephant that would not lower a pillar of wood into
a hole containing a sleeping dog until the dog was chased away (Shoshani
and Eisenberg 1992). Many accounts are available from the wild, too. For
example, the respected naturalist Tony Archer (personal communication)
witnessed a young male repeatedly tusk a crippled female who had fallen
behind her family. A large, older female (possibly a relative) suddenly ran
from 40 meters away to chase the male and, having seen him off, returned
to touch the other female gently on her withered leg. The specific behav-
ior of the older female also indicated that she might have understood the
particular feelings of the crippled female, possibly demonstrating her pos-
session of theory of mind.
While it may be possible to explain some of these accounts as examples
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of genetically wired behavior or trial-and-error learning, without invoking
insight or an ability of elephants to understand that others see, feel, and
know (i.e., theory of mind) recent work by geneticist Moti Nissani (2004)
indicates we should be cautious in drawing such conclusions. In three ex-
periments using captive individuals, Wanda and Winky, Nissani explored
the question of whether elephants are capable of theory of mind and in-
sight, by focusing on whether elephants know that people use the sense of
sight (see Varner, Chapter 3, for more details). Although Nissani’s research
is not conclusive, it adds to growing evidence supporting the view that ele-
phants may be capable of both insight and theory of mind.
In the human brain, there exist multiple mirror neuron systems. Mir-
ror neurons are cells in the brain that specialize in carrying out and un-
derstanding not only the actions of others but their intentions as well as the
social meaning of complex behavior and secondary emotions such as dis-
gust, shame, and guilt. By simulating a “mirror image” of another’s actions
in our brain we are able to empathize, imitate, and acquire language. Mir-
ror neurons are, in effect, the source of our theory of mind. In a sense, mir-
ror neurons absorb culture directly, with each generation teaching the next
through sharing, imitation, and observation (cited in Blakeslee 2006).
Research by neuroscientist Giacomo Rizzolatti, on mirror neurons in
the brains of monkeys have found that these cells, too, are active in situa-
tions that would inspire empathetic feelings (described in Blakeslee 2006).
Some elephants have been observed attempting to raise an immobilized
or dying elephant to its feet (J. Poole and C. Moss, personal observation;
Douglas-Hamilton et al. 2006) or returning to the corpse of a dead com-
panion many times over the course of days and weeks (Douglas-Hamilton
et al. 2006). Taken together, these mirror neuron studies and elephant ob-
servations suggest that monkeys and probably other highly social species
like apes, elephants, dolphins, and even dogs experience empathetic feel-
ings and that the basis for theory of mind exists in a network of cells with
the same origin as our own more complex capability.
Some years ago one of us (J. Poole) watched elephant Eliot flinch vio-
lently as Eudora, 10 meters away, reached out her trunk to test if an electric
fence was connected. We suggest that Eliot’s mirror neurons were actively
firing as she experienced a moment of fear in anticipation of Eudora’s po-
tential shock. Vocal production learning and other imitative abilities in ele-
phants may be yet another piece of evidence in the jigsaw puzzle pointing
to elephant theory of mind.
Self-Recognition in Elephants
Self-directed behaviors in front of a mirror have been interpreted as indic -
ative of self-recognition (Gallup 1970). Furthermore, researchers maintain
that if an animal recognizes itself, then it must have, however rudimentary,
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a sense of self. Mirror self-recognition (MSR) has, thus, been expected to
correlate with higher forms of both altruistic behavior and empathy.
Dozens of primate species have been exhaustively tested for self-recogni-
tion using the mirror-mark test, and only the great apes have conclusively
passed the test (Gallup, Anderson, and Shillito 2002). In 2001, mirror self-
recognition was demonstrated in bottlenose dolphins, a species also noted
for complex social behavior and empathetic traits (Reiss and Marino
2001). Recently, Plotnik et al. (2006) demonstrated that elephants, too, are
capable of mirror self-recognition (see also Varner, Chapter 3). Plotnik et al.
postulate that MSR in apes, dolphins and elephants offers persuasive evi-
dence for convergent evolution in the self-other distinction that underlies
the social complexity, cooperation and altruistic tendencies noted among
these large-brained mammals.
Reaction to Trauma and Death
Recent work by trauma researchers Bradshaw et al. (2005) suggests that early
disruption of attachment can result in social trauma that may affect the phys-
iology, behavior, and culture of elephants over generations. Such disruption
occurs during culls where adults are killed and infants are spared during cap-
ture and abusive training of infants for captive use; and during high-level
poaching. An intact social order may buffer trauma, but as human popula-
tions increase more elephants are living in environments influenced by re-
lentless human disturbances. The consequences of these experiences can be
seen in abnormal levels of aggression in the wild (Slotow et al. 2000) and in
captivity (see also Hancocks, Chapter 13 in this volume).
A discussion of elephant cognitive capacities would be incomplete
without the mention of elephants’ reaction to death. Elephants exhibit a va-
riety of responses to dying or dead elephants and their bones, including
touching with the trunk and feet, attempted lifting and carrying of the
body or bones, mounting, feeding, body guarding, covering, and burying
(Douglas-Hamilton and Douglas-Hamilton 1975; Moss 1975, 1988, 1992;
BBC Natural History Unit 1992, 2005; Poole 1996; Payne 2003; Douglas-
Hamilton et al. 2006; McComb et al. 2006). Although elephants exhibit this
behavior primarily toward other elephants, they may also stand over or
cover the bodies of humans or other animals that they kill (Poole 1996) or
of those that they find killed by predators (N. Njiraini, S. Sayialel, and N.
Sayialel, personal communication). People have known about the reaction
of elephants to death for thousands of years (Meredith 2001) and as far as
we are aware no one has yet come up with a plausible alternative to the ex-
planation that elephants have, however basic, a concept of death.
In summary, relative to other mammals, including humans, elephants are
unusually long-lived and exhibit a high degree of social complexity. Their
91 ELEPHANT SOCIALITY AND COMPLEXITY
—volume).
[FIGURE] Figure
4.6 about here
1.Wemmer,Elephants and Ethics 2/1/08 2:23 PM Page 91

development includes social learning and behavioral innovation, both of
which are manifested in the use and modification of rudimentary tools and
in vocal learning. Elephants have extensive neocortical development, very
good memory and are evidently adept users of Machiavellian intelligence.
Mirror self-recognition by elephants indicates self-awareness (Plotnik et al.
2006), and numerous observations suggest elephants have a rudimentary
theory of mind and anticipatory planning capabilities that may include
imagining future events, such as pain inflicted on themselves and others
and possibly their own deaths. Although many other species may rival ele-
phants in one capability or another, there are few that equal or surpass ele-
phants in the totality of their social and behavioral complexity.
The evidence presented in this chapter, including the recently published
Bradshaw et al. 2005 report on the long-term effects of trauma, indicates
that we must err on the side of caution when welfare issues are being
weighed, and that individual elephant well-being, not commercial gain,
must be the priority. This conclusion has broad implications for the man-
agement of elephants in the wild and in captivity. It is not within the scope
of this essay to set guidelines for the treatment and care of elephants, but
certain guiding principles, most of which are discussed in Clubb and Ma-
son’s report about European zoo elephants (2002) and in Kane, Forthman,
and Hancocks (2005a, 2005b), should be put into practice.
92 OVERVIEW OF ELEPHANT PHILOSOPHY AND SCIENCE
Figure 4.6 Early disruption of attachment such as occurred during culls where
adults were killed and infants were spared can result in social trauma that may
affect the physiology, behavior, and culture of elephants over generations.
Photograph courtesy of Oria Douglas-Hamilton
1.Wemmer,Elephants and Ethics 2/1/08 2:23 PM Page 92

As large, highly social, and intelligent animals, elephants require ample,
environmentally complex space and a sufficient number of conspecifics for
social contact and learning. We should be moving toward a situation in
which institutions are permitted to keep elephants only if the captive situ-
ations provide sufficient space for adequate exercise and stimulation and
allow the elephants to choose among social partners.
In seasonally cold climates, elephants are usually restricted to indoor
barns when the outside temperature goes below about 4.4 degrees centi-
grade. In zoos that experience long, severe winters elephants may be in-
doors, with inadequate space, exercise, and social stimulation for months
at a stretch. Because of the many negative consequences of restricted space
on elephant physical and emotional well-being, zoos located in areas with
prolonged cold winters should be precluded from keeping elephants.
Elephants (including males) should be allowed access to a social group,
not kept in isolation. Males should be allowed to remain with their family
until the age of natural dispersal; females should remain together for life.
During parturition females should remain unchained and in the company
of family members, particularly if experienced females are present. Infants
and calves should not be removed from the care of their mothers and fam-
ily members. Most elephant mothers in zoos were not themselves exposed
to allomothering experiences, and their very poor success as mothers is a
clear consequence. The tradition of removing an elephant from its social
group for the purpose of exchange with other zoos or circuses should cease.
All forms of physical discipline and punishment must be discontinued and
chaining should stop unless absolutely necessary for veterinary care. Thus,
on the basis of all these criteria, circuses are not an appropriate environ-
ment for elephants and most zoos will have to make substantial changes to
meet the most basic needs of elephants.
In the wild the practice of abducting young elephants from their fami-
lies should end. The culling of elephants should be avoided except where
all other options have been exhausted. When culling is deemed essential it
should include whole families; infants and calves should not be spared for
export to zoos, circuses, safari parks, or private reserves. The parallel prac-
tice of introducing traumatized youngsters to new areas without adult role
models should stop. Alternative practices to culling such as translocation
and birth control also have welfare implications, and these must be care-
fully evaluated. Human-elephant conflict is the cause for increasing ethi-
cal dilemmas, and in cases where it is deemed that an elephant must be eu-
thanized, it should be done efficiently and humanely.
What is the right way to treat beings such as elephants? As writer
Douglas Chadwick (1992, 475) states, “If a continuum exists between us
and such beings in terms of anatomy, physiology, social behavior, and in-
telligence, it follows that there should be some continuum of moral stan-
93 ELEPHANT SOCIALITY AND COMPLEXITY
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dards.” Based on available evidence, the time has come for us to move be-
yond old patterns in the treatment of elephants. We must acknowledge and
accept new standards for the future.
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CONTRIBUTORS
469
Lori Alward, PhD
Tacoma Community House
Tacoma, Washington, USA
Joseph C. E. Barber, PhD
Disney’s Animal Kingdom
Orlando, Florida, USA
Janine L. Brown, PhD
Smithsonian’s National
Zoological Park
Conservation
and Research Center
Front Royal, Virginia, USA
Jacob V. Cheeran, DVM
Project Elephant
Government of India
Thrissur, India
Catherine A. Christen, PhD
Smithsonian’s National
Zoological Park Conservation
and Research Center
Front Royal, Virginia, USA
Smithsonian Institution
Archives
Washington, DC, USA
Rosaleen Duffy, PhD
Professor of International Politics
University of Manchester
Manchester, United Kingdom
Yudha Fahrimal, DVM
Universitas Syiah Kuala
Darussalam, Banda Aceh
Sumatra, Indonesia
Richard Fayrer-Hosken,
BVSc, PhD
Department of Large Animal
Medicine
College of Veterinary Medicine
University of Georgia
Athens, Georgia, USA
Marie Galloway
Smithsonian’s National
Zoological Park
Washington, DC, USA
Marion E. Garaï, PhD
Space for Elephants Foundation
KwaZulu Natal, South Africa
Jane Garrison
Los Angeles, California, USA
Hank Hammatt
Elephant Care International
Hohenwald, Tennessee, USA
David Hancocks
Melbourne, Australia
Rebecca Hardin, PhD
Department of Anthropology
and School of Natural Re-
sources and Environment
University of Michigan
Ann Arbor, Michigan, USA
Harvard Academy for Interna-
tional and Area Studies
Cambridge, Massachusetts, USA
Michael Hutchins, PhD
The Wildlife Society
Bethesda, MD, USA
Graduate Program in
Conservation Biology and
Sustainable Development
University of Maryland
College Park
Maryland, USA
Center for Conservation and
Behavior
Georgia Institute of Technology
Atlanta, Georgia, USA
Mike Keele
Oregon Zoo
Portland, Oregon, USA
Winnie Kiiru, MSc
Born Free Foundation
Nairobi, Kenya
Michael D. Kreger, PhD
U.S. Fish and Wildlife Service
Arlington, Virginia, USA
Fred Kurt, PhD
Research Institute of Wildlife
Ecology
University of Veterinary
Medicine
Vienna, Austria
1.Wemmer,Elephants and Ethics 2/1/08 2:26 PM Page 469

Dhriti K. Lahiri Choudhury, PhD
Department of English
Rabindra Bharati University
Calcutta, India
John Lehnhardt
Disney’s Animal Kingdom
Orlando, Florida, USA
Khyne U Mar, PhD, FRCVS
Elephant Family-UK
London, United Kingdom
Jill D. Mellen, PhD
Disney’s Animal Kingdom
Orlando, Florida, USA
Susan K. Mikota, DVM
Elephant Care International
Hohenwald, Tennessee, USA
Gary W. Miller
Disney’s Animal Kingdom
Orlando, Florida, USA
Cynthia J. Moss, ScD
Amboseli Elephant Research
Project
Amboseli Trust for Elephants
Nairobi, Kenya
Joyce H. Poole, PhD
Amboseli Elephant Research
Project
Amboseli Trust for Elephants
Nairobi, Kenya
470 CONTRIBUTORS
Raman Sukumar, PhD
Center for Ecological Sciences
Indian Institute of Science
Bangalore, India
Gary Varner, PhD
Philosophy Department
Texas A&M University
College Station, Texas, USA
Christen Wemmer, PhD
Smithsonian’s National
Zoological Park
Conservation and
Research Center
Front Royal, Virginia, USA
California Academy of Sciences
San Francisco, California, USA
Ian Whyte, PhD
South African National Parks
Kruger National Park, South
Africa
Nadja Wielebnowski, PhD
Chicago Zoological Society
Brookfield Zoo
Brookfield, Illinois, USA
Wemmer and Christen/
Elephants and Ethics
Greg D. Rossel
Motorola Corporation
Fort Worth, Texas, USA
Nigel Rothfels, PhD
College of Letters and Science
University of Wisconsin–
Milwaukee
Milwaukee, Wisconsin, USA
Dennis Schmitt, DVM, PhD
Ringling Bros. and Barnum &
Bailey
Department of Veterinary
Services
Polk City, Florida, USA
Department of Agriculture
Missouri State University
Springfield, Missouri, USA
John Seidensticker, PhD
Smithsonian’s National
Zoological Park
Washington, DC, USA
Lalith Seneviratne
Colombo, Sri Lanka
Brandie Smith, MS
Department of Conservation
and Science
Association of Zoos and
Aquariums
Silver Spring, Maryland, USA
1.Wemmer,Elephants and Ethics 2/1/08 2:26 PM Page 470