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Edited by Patricia C. Wright, Elwyn L. Simons, and Sharon Gursky

Rutgers University Press, 2003


Tarsiiformes, or tarsiers for short, are a group of living species whose 
combination of derived and ancient characteristics make them pivotal to 
understanding the roots of primate evolution. These small-bodied, 
nocturnal, solitary creatures resemble lower primates in their behavior and 
morphology, while some genetic evidence aligns them more closely with 
higher primates, such as monkeys, apes, and humans. This unique combination 
of behavior and anatomy makes the tarsier an especially interesting and 
controversial animal for study among primate behaviorists, evolutionists, 
and taxonomists, who view the tarsiers as "living fossils" that link past 
and present, lower and higher primates in the long chain of evolutionary 

Contributors to this volume draw on a range of scientific disciplines to 
provide a detailed examination of the past, present, and future of these 
intriguing primates.

Patricia C. Wright is a professor of anthropology at SUNY, Stony Brook and 
director of the Institute for the Conservation of Tropical Environments, 
Stony Brook and Madagascar. Elwyn L. Simons is James B. Duke Professor of 
Biological Anthropology and Anatomy and the head of the Division of Fossil 
Primates of the Duke University Primate Center where, with Wright, he 
established one of the world's only viable captive tarsier colonies. A 
member of the National Academy of Science (U.S.A.), he is the author of 
Primate Evolution: An Introduction to Man s Place in Nature. Sharon Gursly 
is an assistant professor of anthropology at Texas A&M University.


List of Figures    ix
List of Tables    xiii
Acknowledgments    xv

Patricia C. Wright, Elwyn L. Simons, and Sharon Gursky    1

PART 1: PAST: Origins, Phylogeny, Anatomy, and Genetics

CHAPTER 1: The Fossil Record of Tarsier Evolution
Elwyn L. Simons    9

CHAPTER 2: The Evolution of the Tarsiid Niche
Nina G. Jablonski    35

CHAPTER 3: How Close Are the Similarities between Tarsius and Other
Jeffrey H. Schwartz    50

CHAPTER 4: Morphometrics, Functional Anatomy, and the Biomechanics of 
Locomotion among Tarsiers
Robert L. Anemone and Brett A. Nachman    97

CHAPTER 5: The Axial Skeleton of Primates: How Does Genus
Tarsius Fit?
Friderun A. Ankel-Simons and Cornelia Simons    121

CRAPTER 6: Phylogenetic Position of Tarsiers within the Order Primates: 
Evidence from y-GIobin DNA Sequences
Carla M. Meireles, John Czelusniak, Scott L. Page, Derek E. Wildman,
and Morris Goodman    145

CHAPTER 7: The Phylogenetic Position of Genus Tarsius: Whose Side Are You On?
Anne D. Yoder    161

PART 2: PRESENT: Taxonomy, Behavioral Ecology, and Vocalizations

CHAPTER 8: The Tarsiers of Sulawesi
Colin P. Groves    179

CHAPTER 9: Outline of the Vocal Behavior of Tarsius spectrum: Call 
Features, Associated Behaviors, and Biological Functions
Alexandra Nietsch    196

CHAPTER10: Territoriality in the Spectral Tarsier, Tarsius spectrum
Sharon Gursky    221

CHAPTER11: The Natural History of the Philippine Tarsier
(Tarsius syrichta)
Marian Dagosto, Daniel L. Gebo, and Cynthia N. Dolino    237

CHAPTER12: Can We Predict Seasonal Behavior and Social Organization from 
Sexual Dimorphism and Testes Measurements?
Patricia C. Wright, Sharon T. Pochron, David H. Haring, and Elwyn L. 
Simons    260

PART3: FUTURE: Conservation

CHAPTER13: History of Captive Conservation of Tarsiers
Helena M. Fitch-Snyder    277

CHAPTER14: Are Tarsiers Silently Leaping into Extinction?
Patricia C. Wright    296

The Editors and Contributors    309
Index    313


Why Tarsiers Interest Us

Today tarsiers are relict primates, with only 4-7 species living on a few 
islands in Southeast Asia: although they are small-bodied and nocturnal, 
tarsiers have provided a century of controversy. The combination of derived 
and ancient characters seen in tarsiers makes them pivotal to understanding 
the roots of primate evolution. The tarsiers' unusual diet and reproduction 
provide a unique perspective to understanding primate behavioral ecology, 
while their specialized anatomy shows us how primates can solve a wide 
range of functional needs.

Tarsiers provide one astounding fact after another. The world's most 
carnivorous primate, tarsiers exclusively eat live animals, predominantly 
insects, reptiles, and amphibians (Niemitz, 1984a; Gursky, 2000a). Among 
the smallest primates (80-150 g), tarsier eyes are bigger than their brains 
(Sprankel, 1965). Small and smooth, tarsier brains resemble carnivore 
rather than primate brains (Rosa et al.,1996). Estrous females have swollen 
red vulvas like Old World monkeys (Wright et al., 1986a), and tarsiers 
produce huge newborn infants, up to 25-30 percent of the mother's weight 
(Haring and Wright, 1989). Tarsiers gestate for six months-several weeks 
longer than macaques or capuchins, which are orders of magnitude larger 
than tarsiers (Izard et al., 1985; Gursky, 1997)-yet tarsier mothers wean 
infants within two months of birth-instead of 12-18 months like macaques 
and capuchins (Haring and Wright, 1989). Tarsier mothers give birth to one 
offspring at a time yet have 4-6 nipples (Wright et al., 1986b). Unlike 
other primates that produce large infants, tarsier fathers provide 
relatively little paternal care (Gursky,2000b). Since infants are so heavy, 
mothers and other group members rarely transport them. Instead, a mother 
will park her infant on branches while she forages nearby (Gursky, 2000b). 
Tarsiers can turn their head 180 degrees in either direction (Ankel-Simons, 
2000) enabling them to see both prey and predators. Owls can do this, but 
no other mammal can. Tarsiers have extremely long legs, and large hands and 
feet for their body size-adaptations for leaps of Olympic dimensions 
(Fleagle, 1998). Their anklebones are elongated-up to four times longer 
than other same-sized primates-and their leg bones (fibula and tibia) are 
fused, the only primate to have this special anatomy for extraordinary 
leaping (Woollard, 1925; Gebo, 1987).

Unresolved Questions

The relationship of tarsiers to other primates, both living and fossil, has 
been a source of debate for over a century. No other primate generates so 
much controversy. What is the tarsier's true phylogeny? Morphological 
evidence, including soft-tissue characteristics, suggests that tarsiers are 
closely related to small nocturnal prosimian primates (femurs, lorises, and 
bush babies), called strepsirhine primates. However, another suite of 
anatomical and reproductive characters suggests that tarsiers may be more 
closely related to monkeys, apes, and humans, called haplorhine primates.

The controversy extends to the fossil record. Are tarsiers derived from 
omomyids, a group of Eocene prosimians? Are they from their own special 
branch? Did they share a common ancestry with the stock that produced 
monkeys, apes, and humans, the adapids? Recently the debate has taken a 
geographic twist: did tarsiers arise in Africa (Simons and Bown, 1985) or 
Asia (Beard, 1998)?

And as for modern tarsiers, how many extant species are there? The answers 
range from three to seven. How do these rarely studied species behave in 
the wild? What is their social organization? And should tarsiers be a 
conservation priority?


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Posted Date:   6/25/2003