The Amboseli Elephants: A Long-Term Perspective on a Long-Lived Mammalby Cynthia J. Moss (Editor), Harvey Croze (Editor), Phyllis C. Lee (Editor)
Elephants have fascinated humans for millennia. Aristotle wrote of them with awe; Hannibal used them in warfare; and John Donne called the elephant “Nature’s greatest masterpiece. . . . The only harmless great thing.” Their ivory has been sought after and treasured in most cultures, and they have delighted zoo and circus audiences worldwide for
Elephants have fascinated humans for millennia. Aristotle wrote of them with awe; Hannibal used them in warfare; and John Donne called the elephant “Nature’s greatest masterpiece. . . . The only harmless great thing.” Their ivory has been sought after and treasured in most cultures, and they have delighted zoo and circus audiences worldwide for centuries. But it wasn’t until the second half of the twentieth century that people started to take an interest in elephants in the wild, and some of the most important studies of these intelligent giants have been conducted at Amboseli National Park in Kenya.
The Amboseli Elephants is the long-awaited summation of what’s been learned from the Amboseli Elephant Research Project (AERP)—the longest continuously running elephant research project in the world. Cynthia J. Moss and Harvey Croze, the founders of the AERP, and Phyllis C. Lee, who has been closely involved with the project since 1982, compile more than three decades of uninterrupted study of over 2,500 individual elephants, from newborn calves to adult bulls to old matriarchs in their 60s. Chapters explore such topics as elephant ecosystems, genetics, communication, social behavior, and reproduction, as well as exciting new developments from the study of elephant minds and cognition. The book closes with a view to the future, making important arguments for the ethical treatment of elephants and suggestions to aid in their conservation.
The most comprehensive account of elephants in their natural environment to date, The Amboseli Elephants will be an invaluable resource for scientists, conservationists, and anyone interested in the lives and loves of these extraordinary creatures.
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The Amboseli ElephantsA Long-Term Perspective on a Long-Lived Mammal
The University of Chicago PressCopyright © 2011 The University of Chicago
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Chapter OneThe Amboseli Elephants: Introduction
Cynthia J. Moss, Harvey Croze, and Phyllis C. Lee
Historical Perspectives on Elephant Studies
Elephants have fascinated humans for millennia. Aristotle wrote of them with awe; people in Asia have tamed, trained, and revered them for centuries; Hannibal used them in warfare; and John Donne called the elephant "Nature's great masterpiece ... the only harmless great thing." Ivory has been sought and treasured in most societies at great cost to elephant populations. Eventually elephants were put on display in zoos and made to perform in circuses where, sadly, they still draw the laughter of children and the jibes of adults. It was only in the second half of the 20th century that people started to take an interest in elephants as elephants, that is, as free-living biological and ecologically functioning beings. This volume aims to provide a portrait of what the Amboseli Elephant Research Project (AERP) has contributed to our current knowledge of elephants as integrated components of their ecosystems and as individuals with potentially long and intricately involved social lives.
People who live in contact with elephants invariably are interested in them, whether they try to hunt them or avoid being killed by them. Knowledge of elephants among central African rainforest pygmies or savannah pastoralists is traditionally colored by myth, magic, and ethno-ecological reality derived from their experience and close contact. The Europeans who arrived in Africa to hunt "big game" had somewhat different attitudes. Those foreign hunters wrote numerous books and articles with titles such as On Killing Elephant (Bell 1931). Most such books were narratives of the writer's fearlessness in the face of massive charging bulls. Very little natural history appeared in their accounts, and the few descriptions of elephant life were often erroneous. Seen through the eyes of 19th-and 20th-century European men, the social life of wild animals was a clear reflection of the beholder's worldview—or perhaps more a reflection of what he wished the world to be. Almost all of them described elephants as living in a harem system with a single herd bull and his many females. Hunters' tales dominated in descriptions of the behavior of Asian elephants; the more knowledgeable mahouts or keepers of captive animals had relatively few opportunities in the colonial context to correct these misapprehensions.
In the late 1950s and early 1960s, a new group descended on Africa to pursue wildlife. These were biologists, ecologists, physiologists, parasitologists, and other scientists, whose technique for studying elephants was to shoot them and examine the carcasses. Much was learned in this way, but the elephants' death limited opportunities to study behavior and to gather longitudinal data. In the 1960s, ethologists who had been taught by Niko Tinbergen at Oxford came out to Africa to study living animals in the wild. Tinbergen, along with Konrad Lorenz and Karl von Frisch, jointly received the Nobel Prize in 1973 for developing ethology—the experimental study of animal behavior under natural conditions whenever possible—as a scientific discipline. One of Tinbergen's students was Iain Douglas-Hamilton, who began a pioneering study of wild elephants in Lake Manyara National Park, Tanzania, in 1965.
Douglas-Hamilton's study (on which Cynthia Moss worked for a year) was based on the ability to recognize individual elephants over time. Each animal in the study population was named or numbered, and its movements, activities, associations, and relationships were recorded over a four-year period. Using this novel approach, a new understanding of elephant social life, behavior, and ecology began to emerge. At the same time, Harvey Croze was working in the Serengeti National Park as a Tinbergenian post-doctoral researcher studying the interactions of elephants and their habitat. In 1972, at the suggestion of David Western, Moss and Croze started the AERP, which stands today as the longest continuously running elephant research project in the world. At the time this introduction is being written, the project is in its 38th year. Neither of the founders thought at the time that they would be personally involved in the project for so many years, but they had planned from the beginning for AERP to be a long-term research commitment. The guiding philosophy was that elephants are long-lived animals and that to gather the data necessary to understand and describe their ecology and behavior would need a lifetime—both elephant and human
Background to the Study Population
Forty years ago, the Amboseli elephant population was known to be relatively small, to co-exist with Maasai pastoralists in a semi-arid savannah (the dynamics of Amboseli's ecology are detailed in chapter 2), and to be both approachable and observable (Western 1973). The big males were being sport hunted, and there were low levels of poaching and conflicts with people, but inside the protected area, they were tolerant of vehicles. They appeared to be good candidates for study.
During the 1970s and 1980s, the African continent saw a catastrophic decline in its elephant populations. In almost every country where elephants ranged, ivory poaching drastically reduced numbers. Throughout that period of changing fortunes for Africa's elephants, only a few populations, such as that in Amboseli, were relatively well protected. Three circumstances in the Amboseli ecosystem contribute to its relatively undisturbed state. The land surrounding the National Park belongs to the Maasai people, who traditionally have been tolerant of wildlife and at the same time inhospitable to outsiders hunting or poaching on their land. In addition, the Amboseli elephants have never been culled as part of a park management program nor have they been fenced into a protected area. Finally, the research project itself has provided wildlife and anti-poaching authorities with additional eyes and ears and a nearly full-time vigilante presence in the field.
In the comparative absence of poaching and culling (but see box 19.1), the Amboseli elephants have increased slowly in numbers since the late 1970s. The population in 2008 at just over 1,500 remains relatively small but is nonetheless very important for Kenya and the rest of Africa. Amboseli is one of few places in Africa where the elephant age structure has not been drastically skewed by past poaching. Known individuals span the range from newborn calves to old matriarchs in their 60s and—exceptional these days—large adult bulls in their 50s and early 60s. With a natural age structure and intact social organization, the Amboseli elephant population has become increasingly important as a source of baseline data on elephant social and reproductive behavior and population dynamics. As such, it is used as a model for determining the status of other elephant populations in Africa and Asia and as a yardstick for assessing the extremes that captive elephants experience.
Threads and Themes
Four underlying themes run throughout the chapters in this volume: longevity, size, intelligence, and the future of the elephants in a changing and threatened ecosystem. The book is divided into five parts, which are explained in detail below (See annex 1.1 for a glossary of Maa place-names and box 1.1 for definitions of common terms used throughout the book).
Longevity in the Amboseli context has two facets: the elephants and the study itself. The commitment of a group of cooperating but highly individualistic researchers, some of whom have collaborated for four decades, is almost as remarkable as the unfolding of an animal's lifespan over nearly seven decades.
The Amboseli study presents an important opportunity to explore longevity in a natural evolutionary and functional context. Longevity is not easily studied in natural populations; only studies of a few large mammals such as lions (Mosser and Packer 2009), red deer (e.g., Kruuk et al. 1999b), and sheep (e.g., Festa-Bianchet, Jorgenson, and Réale 2000) have provided longitudinal individual life history data. Behavioral aging—changes in behavior or functions with age—is a poorly understood attribute common among all long-lived animals (Finch 1994). Some individuals age rapidly while others resist the effects of age. Most species that have been used as models for evolutionary and behavioral aging are small and short-lived, so they can be reasonably observed and manipulated over the duration of a research grant. In this volume, we add elephants to these classic studies, but we should stress that despite the unique dataset analyzed and presented in the chapters that follow, we have still witnessed only half of an elephant's potential lifespan. While we have been careful not to presume that we can as yet accurately assess genetic fitness over a full lifespan, we do explore the individual fitness consequences of much of the behavior.
Size, of both the animal and its spatial requirements, is the second theme. The elephant is at the extreme end of the scale in terms of terrestrial mammalian body mass and energetic relationships. It also has to cope both individually and socially with the heterogeneity and dimensions of a habitat that ranges from swamp to dense bushland to bare soil over thousands of square kilometers. Large size and a long growth period have consequences for reproduction and sociality as well as for the retention and manipulation of information about companions, enemies, and the environment.
Thus, the third theme is the exceptional intelligence that provides elephants an evident adaptive edge in habitats that vary over time in unpredictable ways. We address this issue by providing descriptive but data-rich portraits of elephant behavior, gestures, and signals, which are contextualized by experimental manipulations of communication and explorations of social knowledge. We also emphasize throughout these chapters the dynamic nature of elephant sociality: fission-fusion, where each individual is embedded in a knowledge-based network of others. While fission-fusion as a system of aggregation may not require any particular cognitive abilities, as evidenced by fish schools or flocks of roosting starlings (see, for example, Aureli et al. 2008), elephants have to rely on memory, knowledge, and information exchange to maintain their networks, and thus sociality both drives and enables elephant intelligence.
The final theme, which looms like a dreadful portent, is the uncertainty of the elephant population's future in the face of changing social and economic circumstances in the ecosystem surrounding Amboseli National Park. That uncertainty has over the years caused the project to stretch beyond its original research mandate and reach out to the local community, to demonstrate benefits from the elephants and their ecosystem, and to form partnerships with the Maasai. These partnerships provide the project with valuable information about ecosystem occupancy and community concerns and aim to maintain an atmosphere of goodwill toward elephants that is fundamental to their future.
All four themes situate the study and the volume in a single place: Amboseli. This volume is special in that it approaches a single species in one place from many perspectives, exploring how it makes a living in a specific but constantly changing habitat over a 60-plus-year lifespan from birth to death. Previous studies of single ecosystems (e.g., Serengeti [Sinclair and Arcese 1995]) have perforce skimmed the surface of behavior and ecology of many of their constituent species over time, while studies of a single species (e.g., roe deer [Anderson et al. 1998]) or species group (e.g., cetaceans [Mann and Smuts 1998; Mann et al. 2000]) have had to compare populations rather than examine them over time. In the pages that follow, we intentionally keep the perspective focused on the elephants of Amboseli and do not belabor comparisons with studies of all other elephant populations. Such comparisons will emerge in time, we hope, as a result of this volume.
The focus on a single population over time provides a stable reference point for theoretical understanding; being large, long-lived, and distributed across habitats from dense forests to deserts, hemmed in by a variety of human landscapes, means that elephant populations exhibit variation at all levels and scales. The ability to examine variation in context for one population should enhance our ability to compare, contrast, and make sense of the sweep of elephant variation across the species' range.
What the Book Is About
This volume has had the equivalent gestation length of five—almost six—elephant calves. As more researchers joined the project, bringing their individual theoretical and practical backgrounds and expertise, enthusiasm grew for compiling all our knowledge in a single publication. In September 1998, the nominated section editors, hosted by the White Oak Conservation Center in northern Florida, met to discuss content and a workplan. By October 1999, when we met again as a working group to discuss the analysis of long-term datasets at the National Center for Ecological Analysis and Synthesis in Santa Barbara, California, we confirmed the importance of the work and had mapped out a clear way forward for tackling the long-term dataset.
Many readers of this volume will not recall those days at the start of the project in the early 1970s, when we were in the computational Stone Age: we had only just shelved our slide rules and begun trying out the new handheld programmable calculators. Our original data were designed to be compatible with 80-column IBM Hollerith punch cards, and this format was retained consistently across a variety of now-extinct data-entry modes and analysis protocols. Another fortunate decision was to cast the project's location data collection into a 1 km2 map grid, which, although well before the days of affordable commercial satellite data and global positioning systems and at the dawn of geographic information systems, proved most useful for subsequent mapping and spatial analysis (see appendix 1).
The construction, cleaning, and verification of a long-term amalgamated sightings database started in 1996 and continued until 2003 and for this reason, much of the analysis in the volume uses sightings data for only the first 30 years of the project, that is, up to the end of 2002. New data have been constantly added, but the analysis had to have an end point, and 30 years seemed a reasonable cutoff. As chapters have evolved and been revised over time, we have incorporated additional data where necessary; some chapters have included political changes and novel observations right up to the moment of going to press. Analyses of demographic and population data extend until 2006 in order to use recent events to provide further insights.
Thus, much of the data we present have not appeared in previous publications because they are based on a long-term dataset that was constructed and made manageable only during the course of writing the book. Many chapters build upon published peer-reviewed research but take advantage of the whole of the dataset to revisit and reanalyze earlier research questions. The chapters herein are thus a blend of previously published but augmented research merged with entirely novel analyses resulting from research questions mooted over the 10-year gestation of the volume. We have subjected the new analyses to peer review, commentary, and revision.
Throughout the book, we often use statistics to argue a point. But when statistical tests are based on huge sample sizes such as the sightings database (N circa 30,000), statistical power is high but explanatory effect may actually be rather small. We have thus tried to explore biologically sensible effects rather than simply using P values. Despite having nearly 40 years of almost daily observations of slowly reproducing, very long-lived individuals, analysis is further bedeviled by having relatively few records of those rare events that make up the unique life history of any individual. As Dick Byrne observes (see chapter 11), the value of very long-term studies is that as time goes on, the unexpected and, indeed, the previously unknown are eventually seen. We have tried to maintain the "individual" perspective over a lifespan as a focus within each of the themes in the book.
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Meet the Author
Cynthia J. Moss is the director of the Amboseli Trust for Elephants and the author of Elephant Memories: Thirteen Years in the Life of an Elephant Family. Harvey Croze is a trustee for the Amboseli Trust for Elephants and coauthor of Pyramids of Life: An Investigation of Nature’s Fearful Symmetry. Phyllis C. Lee is professor of psychology at the University of Stirling. She is the author or editor of several books, including Threatened Primates of Africa and Comparative Primate Socioecology.
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