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Tupai: A Field Study of Bornean Treeshrews

Louise H. Emmons
Foreword by Harry W. Greene
Copyright Date: 2000
Edition: 1
Pages: 287
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  • Book Info
    Book Description:

    Treeshrews suffer from chronic mistaken identity: they are not shrews, and most are not found in trees. These squirrel-sized, brownish mammals with large, dark, lashless eyes were at one time thought to be primates. Even though most scientists now believe them to belong in their own mammalian order, Scandentia, they still are thought to resemble some of the earliest mammals, which lived alongside the dinosaurs. This book describes the results of the first comparative study of the ecology of treeshrews in the wild. Noted tropical mammalogist Louise H. Emmons conducted this pathbreaking study in the rainforests of Borneo as she tracked and observed six species of treeshrews. Emmons meticulously describes their habitat, diet, nesting habits, home range, activity patterns, social behavior, and many other facets of their lives. She also discusses a particularly interesting aspect of treeshrews: their enigmatic parental care system, which is unique among mammals.

    eISBN: 978-0-520-92504-5
    Subjects: Zoology

Table of Contents

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  1. Front Matter
    (pp. i-iv)
  2. Table of Contents
    (pp. v-vi)
  3. List of Illustrations
    (pp. vii-x)
  4. List of Tables
    (pp. xi-xii)
  5. Foreword
    (pp. xiii-xvi)
    Harry W. Greene

    Tupai: A Field Study of Bornem Treeshrewsis the second volume in the University of California Press’s new series on organisms and environments. Our main themes are the diversity of plants and animals, the ways in which they interact with each other and with their surroundings, and the broader implications of those relationships for science and society. We seek books that promote unusual, even unexpected connections among seemingly disparate topics; we want to encourage writing that is special by virtue of the unique perspectives and talents of its authors. Our first installment, by Carl E. Bock and Jane H. Bock,...

  6. Acknowledgments
    (pp. xvii-xviii)
  7. CHAPTER 1 Tupai: An Introduction
    (pp. 1-6)

    Treeshrews¹ suffer from chronic mistaken identity: first, they are not shrews; second, most are not found in trees; and third, what they really are (among mammalian orders) has never been agreed on. The first treeshrew recorded by a Western naturalist was collected in 1780 by William Ellis, surgeon to Captain Cook’s expedition (Lyon 1913). He thought it was a squirrel. Indeed, at first glance, treeshrews are so much like squirrels that people who live where they occur often confuse the two, and they are known by the same common name,tupai,in the MalayIIndonesian language. This local name was the...

  8. CHAPTER 2 The Study Species
    (pp. 7-23)

    Every organism is captive within the architecture of its inherited physical and chemical structure. Its interactions with the outside world (ecology) are limited by its evolutionary constraints. To establish the physical framework from within which treeshrews must function, in this chapter I review selected aspects of their anatomy and physiology that are especially pertinent to their ecology.

    There are two subfamilies of treeshrews, the Ptilocercinae, including only the nocturnal pentail treeshrew (Ptilocerucus lowii), and the Tupaiinae, including the diurnal genera Ananthana, Tupaia, Urogale, and Dendrogale. The members of these two subfamilies are anatomically very unlike, with many of their differences...

  9. CHAPTER 3 The Milieu: Field Study Sites and Habitats
    (pp. 24-37)

    One of two Malaysian states on Borneo, Sabah is a quiet, rural region that occupies the island’s northern tip, sandwiched between the South China and Sulu seas (fig. 3.1). Borneo was once totally-forested, except above the treeline on mountains, but much of its surface is now agricultural, silvicultural, or heavily logged. The sparse human population of Sabah has put relatively little pressure on the land, and the large tracts of forest that remain hold some of the last, best populations of Sumatran rhinos, proboscis monkeys, and orangutans. Beside these well-publicized species lives a rich fauna of little-known smaller mammals (Payne,...

  10. CHAPTER 4 Treeshrews in Their Habitat
    (pp. 38-52)

    The first task at the start of any field study is to find the study species. This can be the most difficult part of fieldwork, and projects sometimes founder at this initial point. Among the most satisfying moments to a field biologist is the instant of realization that one can predict where an animal (or plant) can be found. At that moment one has begun to understand the organism. This account of the results of fieldwork on treeshrews starts at its logical beginning and first field question: where are the treeshrews? I answered this simply, by learning to recognize...

  11. CHAPTER 5 Diet and Foraging Behavior
    (pp. 53-90)

    Its diet is the most fundamental ecological attribute of an animal. Virtually every feature of a species’ biology, from morphology to life history, is linked to what it eats and where, when and how nutrition is acquired. The diet of treeshrews was from earliest reports correctly known to consist of fruit and insects (Cantor 1846). However, this knowledge alone does not distinguish treeshrews from other rainforest mammals, almost all of which also eat fruit and insects (Emmons 1995), nor does it define the ecology of individual species. Below I describe the nature of fruit-and insect-eating by treeshrews, along with the...

  12. CHAPTER 6 Nesting Behavior
    (pp. 91-109)

    Because half of a treeshrew’s time is spent inactive, its nest site and the nest itself are primary defenses against exposure and predation. The absentee maternal care system of treeshrews is linked to their nesting behavior as young are placed in one nest and their mother uses another (Martin 1968). To study the maternal care system, I therefore needed to try to find nests of both adults and young, and learning where treeshrews nested was a pivotal element of the field study. An obvious question about the absentee system is whether treeshrew nests are placed where predation risks can be...

  13. CHAPTER 7 Activity Patterns
    (pp. 110-123)

    Even without other knowledge, a description of the activity pattern of an animal reveals much about its life. As in human endeavors, the amount of time animals spend in various activities is a direct reflection of their basic economics. The activity of treeshrews was recorded by following radio-tagged individuals, rain or shine, from before they left the nest to begin their daily activity until they returned to the nest at its end. The results below are compiled from 185 complete daily records (see chapter appendix). Because it was rare to see the animals, the “activity” described here is an analysis...

  14. CHAPTER 8 Use of Space
    (pp. 124-144)

    A home range estimate is built from a map of the points where an animal is known to have been. The points we know are but a tiny sample of the terrain that the animal might use, and a good deal of thought and theoretical discussion has been devoted to divining the best way to estimate the true home range from isolated points (e.g., the six models summarized in Kenward 1987). Researchers who take radiolocations only every few hours or days do not know how the animal traveled between points, and models that fill in empty spaces, such as “the...

  15. CHAPTER 9 Social Organization
    (pp. 145-168)

    Social organization can be broadly defined as the pattern of interactions between individuals. This pattern determines ecological questions, such as how the physical space of the habitat is used by different classes of individuals and how resources are allocated among them; and also more behavioral and evolutionary questions, such as which individuals produce offspring and what happens to those offspring as they mature. Because treeshrews could not often be watched, I had to infer their social relationships from the deployment of their home ranges and the relative movements of radio-tagged and trapped individuals. The key question of whether treeshrew home...

  16. CHAPTER 10 Life History
    (pp. 169-201)

    One of the most remarkable features ofTupaiabiology is the “absentee” maternal care system. Martin (1968) first described this behavior in captiveT. glis belangeriand then in captiveT. minorandT. tana(D’-Souza and Martin 1974). The basic features of the system Martin (1968) described are (1) the mother gives birth to her young in a nest that she never shares with them, where they stay until weaning; (2) the mother visits the young to nurse them only once every other day for less than five minutes; and (3) the mother does not show any of the...

  17. CHAPTER 11 Predation, Predators, and Alarm Behaviors
    (pp. 202-209)

    The chief antipredator strategy of treeshrews is to avoid being noticed. They do this with silent, inconspicuous movements and by keeping undercover, helped by their cryptic coloration. Little escapes their acute senses and extreme alertness. When a treeshrew spots a suspicious but not immediately dangerous situation, such as a motionless scientist, it is likely to give alarm calls from a discreet distance and safe site, such as a log, brush pile, or vine tangle. If the same threat is moving along, the treeshrew is unlikely to call but freezes watchfully or vanishes silently. Pentail treeshrews sometimes dashed around tree trunks...

  18. CHAPTER 12 Synthesis
    (pp. 210-226)

    The six species of treeshrews each use the rainforest environment in a unique way (fig. 12.1). Three species each segregate sharply from all the others by major differences in activity phase (P. lowii), foraging height (P. lowii, T. minor), or habitat specialization (T. montana). The three others (T. gracilis, T. longipes, T. tana), which coinhabit the lowland forest floor, differ in body size, foraging substrate, and invertebrate prey, as well as in nest sites.

    Species of the two pairs that are most similar in body size,T. minorandT. gracilisandT. montanaandT. longipes,are completely separated by...

  19. APPENDIX I Methods
    (pp. 227-231)
  20. APPENDIX II Fruit Species Collected at Danum Valley
    (pp. 232-236)
  21. APPENDIX III Mammal Species Found on the Study Plots
    (pp. 237-239)
  22. APPENDIX IV Invertebrates in Treeshrew Diets
    (pp. 240-243)
  23. APPENDIX V Consumers of Fruit Species
    (pp. 244-246)
  24. APPENDIX VI Response of Murid Rodents to the Masting Phenomenon of 1990-1991
    (pp. 247-250)
  25. Bibliography
    (pp. 251-260)
  26. Index
    (pp. 261-269)
  27. Back Matter
    (pp. 270-272)