|Elephants are socially complex|
Elephants are well-known for their intelligence, close family ties and social complexity, and they remember for years other individuals and places. They live in a fluid fission-fusion society with relationships radiating out from the mother-offspring bond through families, bond groups, clans, independent males and beyond to strangers.
Special relationships between individual elephants may last a lifetime, though the quality of these relationships and the structure and degree of cohesion in an individual's social network may change through time.
Like many sexually dimorphic mammals, adult male and female elephants live in very different social worlds. A complex network of bonds between individuals and families characterizes the lives of females and their offspring, while fluctuating sexual cycles distinguish the dynamic activities and relationships of males.
Personalities and relationships
Elephants have strong individual personalities that affect how they interact with other elephants, how others perceive them, and how well they are able to influence members of their group. For example, some elephants are popular while others are not. Some elephants show strong leadership qualities, others do not; some are highly social "extroverts", while others are less social, "introverts." In order to better understand the personalities of individual elephants, members of the Amboseli Elephant Research Project (AERP) completed a Personality Questionnaire form (developed for people) for the adult members of Echo's family. Each elephant was rated on a scale of 1-7 on characteristics such as: active, aggressive, apprehensive, confident, curious, deferential, eccentric and so on. You can read about the result in the book by the Amboseli Elephant Research Project, The Amboseli Elephants: A Long-Term Perspective on a Long-Lived Mammal.
No individual or individual personality has more impact on family structure and fortunes than the matriarch, or female leader, of an elephant family. Matriarchs are generally the oldest and largest adult female member of a family. In some cases it is very obvious who the leader of a family is - she is the individual to whom others orient, whose sudden change in activity they note, and whose leadership they turn to in a moment of crisis. In other families, the matriarch's leadership is less clear, and in still others there may be conflict over who is the leader. The reasons for these differences have to do with personality, genetic relationships and history.
Matriarchs express their dominance in both competitive and cooperative situations. Matriarchs have been traditionally thought of as rather autocratic, making most decisions and leading a rather docile group of "followers." While this may be the case in some populations and in some families, it has not been our observation. In general, we find that any member of a family may make a suggestion about a plan of action. Suggestions are typically made by adults, though on occasion even a juvenile may offer a suggestion. Some elephants are natural born leaders and they begin to display their abilities at an early age. The most successful leaders seem to be confident, well-connected individuals who are able to command the respect of others both through their wisdom and charisma. A suggested plan of action by an individual may be ignored, discussed, negotiated, or simply followed without further ado. It is true that if a matriarch makes a suggestion there tends to be less discussion than if a suggestion is made by another member of the family - but even this does not always hold true. Very often the matriarch has to argue her case and in many instances she loses her argument to the combined will of other members of the family (if you want to know how elephants discuss, then you'll have to learn more on the vocalization database!).
So, if the matriarch is not crucial in day-to-day decision-making, when is she important? Joyce believes that successful matriarchs are not self-appointed leaders of their family; they are leaders because members of their family respect them, and they are respected because they have proven over the years that they can be trusted to make wise decisions in a time of crisis. Through the years older females become "repositories" of social and ecological knowledge. So natural leadership qualities (personality) and long experience combined are the makings of a wise matriarch. Individuals living in families with older, more experienced matriarchs seem to gain more (in terms of survivorship) from her experience. However, not all matriarchs make wise decisions. Many families split up soon after the death of a matriarch and Joyce believes that this may be partly related to conflict within the group over the new leader.
What are the qualities of a good matriarch? Why are some matriarchs able to hold their family together while under the leadership of others families fall apart? We do not have answers to these questions though we will hazard an educated guess. A successful matriarch needs to be both genetically and socially well connected to all members of her family. In essence this means that she must use her social skills to show compassion and care toward all members of her extended family (not just her own daughters). She must prove to the others that she is worthy of being their leader - by her display of courage and wisdom in times of crisis, by her awesome memory of places and individuals in tough or dangerous times, by her intricate use of tactics in socially difficult situations, and through her excellent social skills to regularly and consistently build, maintain and reinforce the close bonds within her family. In the end, however, when families become very large, and the costs of competition outweigh the benefits of cooperation, even the best matriarch may not be able to hold a family together.
An elephant family consists of one or more usually related adult females and their immature offspring who feed, rest, move and interact in a coordinated manner and have close and friendly ties. Members of a family show extraordinary teamwork and are highly cooperative in group defense, resource acquisition, offspring care, and decision-making. Members of a family greet one another with a special "Greeting Ceremony." While cooperative behavior occurs most often between closely related individuals, kin selection does not explain all such behavior and a proportion of cooperative behavior within families can be explained by reciprocal altruism. The intricate teamwork witnessed between members of an elephant family is to a large extent mediated by a complex suite of vocalizations. Day to day decision-making involves broad participation including consensus building. In Amboseli families contain anywhere from 2 to 16 adult females (that is, females over the age of 10). A matriarch, usually the oldest and most respected female, leads each family. Families may consist of as few as two or as many as 50 or more individuals.
The term "family" or "family unit" has meant different things to different people. Over more than three decades of observations in Amboseli we have witnessed the formation of new families and the dissolution of others. These changes occur over the course of many years as a complex network of relationships is built up or breaks down, and therefore it can be difficult to define at what point in time a particular group of individuals has become or ceased to be a family unit.
Above the level of the family unit a second tier of relationships may exist within what is termed a bond group. Like the family, bond group membership is also usually determined by genetic relatedness. A bond group may include as many as five or more families, and up to 50 or more individuals. Although the ties between individuals across the bond group are weaker than those within a family, bond group members also have close and friendly ties, form alliances against aggressors, assist in the care of another's offspring, defend one another in times of danger and greet one another in a special way.
Bond groups are usually formed as the bonds within a family weaken and families begin to split up along matrilines. In different habitats there exists an optimum family size where the benefits of group living outweigh the costs of competition for resources. As a threshold is reached and the costs begin to outweigh the benefits the family will begin to break up - especially during periods of low food availability. In Amboseli it appears that mothers, daughters and sisters are most likely to stay together, while the fissures tend to divide cousins.The cohesion of different families and bond groups varies significantly and depends upon a number of factors, including personalities, the degree of relatedness and friendship between individuals, the strength of the matriarchs' leadership, historical events such as deaths of important individuals, the type of habitat, and the season.
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