The primates



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CHAPTER 4

THE PRIMATES


CHAPTER OVERVIEW
This chapter introduces students to the study of living, non-human primates. It discusses the basic classification of primates and all of the classes of living primates. It also examines the similarities and differences between non-human primates and humans as well as the earliest primate species which lived during the Eocene and Miocene.

CHAPTER OBJECTIVES
1. Be able to discuss how humans are related to the other primates and the importance of analogies and homologies in determining this relationship.
2. Know the suite traits that are unique to primates.
3. Understand what prosimians are, where they live, and how they are related to other primates.
4. Know the differences between New World and Old World monkeys.
5. Be able to identify and distinguish between the apes.
6. Understand the threats facing endangered primates around the world.
7. Be able to discuss both the similarities and differences between humans and the other primates.
8. Be able to identify and distinguish between the Early Cenozoic primates and the Oligocene anthropoids.
9. Be able to identify and distinguish between the Miocene hominoid species.


CHAPTER OUTLINE



I. Our Place among Primates
A. Definitions of key terms

1. Taxonomy: the assignment of organisms to categories.

2. Hominoidea (hominoids): the superfamily containing humans and apes.

3. Phylogeny: genetic relatedness based on common ancestry.


B. Phylogenetic Classification

1. Organisms are placed in classifications, which are arranged hierarchically according to degree of genetic relatedness.

2. Phylogenetic classification is a descending hierarchy of classifications, from most inclusive to least inclusive.

3. Species are constituted by organisms whose mating produces viable and fertile offspring.



II. Homologies and Analogies
A. Homologies

1. Similarities that organisms share because of common ancestry are called homologies.

2. The presence of homologies is the principal factor in determining how organisms are assigned to taxonomic categories.
B. Analogies

1. Analogies are similarities between species that are the result of similar adaptation to similar selective pressures--analogies are not the result of common ancestry.

2. The process which leads to analogies is called convergent evolution.
III. Primate Tendencies
A. Grasping (precision grip, thumb opposability, nails instead of claws).
B. Smell to Sight (eye placement, brain organization, and color vision all reflect a primate emphasis upon sight over smell--not that some of these features are not common to all primates in the same degree).
C. Nose to Hand (increasing reliance on sense of touch as opposed to muzzle, whiskers for information).
D. Brain Complexity (the brain areas devoted to thought, memory, and association are more elaborate and proportionally larger).
E. Parental Investment (single offspring births combined with longer development periods stemming from neotony).
F. Sociality (strongly associated with parental investment, cooperative social groups are selected for in part because of the needs arising from primate parenting).
IV. Prosimians
A. Prosimians vs. Anthropoids

1. Prosimians and anthropoids constitute the two suborders of primates.

2. 30 million years ago, prosimians were driven from niches by better adapted anthropoids.
B. Lemurs and Tarsiers

1. Most of the remaining prosimians, by far, are lemurs.

2. These live only in Madagascar, which separated from Africa prior to the development of anthropoids.

3. Tarsiers survived in Asia, where there are monkeys, by adapting to night conditions (monkeys are not nocturnal).


V. Anthropoids
A. Vision

1. Evolutionary changes in vision probably occurred in response to the pressures of an arboreal habitat.

2. Binocular, stereoscopic vision and color vision may have been selected due to the improved depth perception it endows (locomotion, catching insects, identifying edible fruits).
B. The arboreal habitat (climbing, feeding) and the increasingly social environment (mutual grooming, tool making) were likely factors in selecting for increased manual tactility.
C. Proportionately larger (than prosimians) brain mass and emphasis on memory and cognition were likely selected for by the social environment.

VI. Monkeys
A. Platyrrhines and Catarrhines

1. There are two anthropoid infraorders: platyrrhines (flat-nosed, New World monkeys) and catarrhines (sharp-nosed, Old World monkeys, hominoids).

2. Unlike hominoids, monkeys’ rear and fore limbs articulate from their bodies as do dogs’.

3. Most monkeys have tails.


B. New World Monkeys

1. New World monkeys’ traits: universally arboreal, some brachiate, some have prehensile tails (among primates, a trait exclusive to the New World).

2. The brachiation of New World monkeys and the brachiation of gibbons constitute an analogy.
C. Old World Monkeys

1. Old World monkeys are both terrestrial and arboreal.

2. Significant distinctions existing between arboreal and terrestrial Old World monkeys include size (arboreal monkeys are smaller than terrestrial monkeys) and sexual dimorphism (terrestrial males are significantly larger and fiercer than terrestrial females, while little or no such differentiation exists among arboreal monkeys).
VII. Apes
A. Old World Monkeys comprise the superfamily Cercopithecoidea, while humans and apes are in the superfamily Hominoidea.
B. Hominoidea is subdivided into three families.

1. Hominids (humans and their fossil ancestors)

2. Pongids (“great apes”: gorillas, chimpanzees, and orangutan)

3. Hylobatids (gibbons and siamangs)

4. Recent biochemical evidence suggests that gorillas and chimpanzees are almost as closely related to humans as they are to each other.
C. Gibbons

1. Gibbons are small, arboreal, mate for life, and produce few offspring.

2. Their principal mode of locomotion is brachiation.
D. Orangutans

1. Orangutans are relatively large (up to 200 pounds), solitary, and markedly sexually dimorphic.

2. Orangutans move between arboreal and terrestrial habitats.
E. Gorillas

1. Gorillas are large (up to 400 pounds), the most sexually dimorphic of all primates, and are primarily terrestrial.

2. They live in relatively stable social groups, typically led by a mature silver-back male.
F. Chimpanzees

1. There are two kinds of chimpanzee: the common (Pan troglodytes) and the pygmy (Pan paniscus).

2. The common chimpanzee is found in western central Africa and western Africa.

3. Size range is up to 200 pounds, and sexual dimorphism is proportionally the same as in humans.

4. Chimpanzee social organization is relatively well-known, because of the longitudinal studies done by Goodall and other primatologists.

G. Bonobos

1. Bonobos belong to the species Pan paniscus.

2. Bonobos live in the humid forests of the Democratic Republic of Congo.

3. Bonobo communities are female-centered, peace-loving, and egalitarian.

4. Bonobos frequently use sex to avoid conflict within the community.
VIII. Endangered Primates
A. Humans are the only hominoids that are not endangered.
B. Deforestation, poaching, and the capture of primates have all contributed to the demise of wild primate populations.
IX. Human-Primate Similarities
A. Learning

1. Neotony and life in cooperative social groups allow primates to learn behavior from their fellows, rather than relying only on genetically encoded behaviors.

2. Learned behavior has been observed in monkeys as well as apes.
B. Tools

1. Tool use allows primates to adapt to a wider range of niches more quickly than physiological adaptation alone (although primates are not the only animals that use tools).

2. Wild chimps have been observed constructing tools.
C. Predation and Hunting

1. Hunting is a regular and normal component of wild chimpanzee behavior.

2. Hunting by chimps is both opportunistic and planned.

3. Wild chimpanzees have been observed hunting consistently, using cooperative techniques, with some sex specialization (males hunt more than females).


D. Aggression and Resources

1. The capacity for hunting exists among many different primates, but expression of this capacity can depend upon environmental pressure and opportunity.

2. Observations of chimps and orangutans indicate that aggressive behavior (“warfare,” in some chimp cases) may increase when territorial encroachment occurs.
X. Human-Primate Differences
A. Sharing, Cooperation, and Division of Labor

1. Sharing and cooperation are common to most primates; however, humans do it much more complexly.

2. Human foraging bands tend to have a sexual division of labor (e.g., men hunt, women gather); other primates do not.

3. Homo sapiens is the only primate species that engages in food sharing consistently on a large scale.


B. Mating and Kinship

1. Human females do not experience estrus.

2. Marriage and kinship are two exclusively, universally human systems that give identity and stability to certain types of human relationships in a way that is absent from other primate social systems.

XI. Chronology
A. The history of vertebrate life on earth is divided into three eras: Paleozoic; Mesozoic; and Cenozoic.

1. Each era is divided into periods.

2. Each period is divided into epochs.
B. Anthropologists are concerned with the Cenozoic era, which includes two periods: Tertiary and Quaternary.

1. The Tertiary period has five epochs: Paleocene; Eocene; Oligocene; Miocene; and Pliocene.

a. A wide range of small mammals, some possibly ancestral to primates, appeared during the Paleocene.

b. Prosimian-like fossils abound in strata dating to the Eocene.

c. The first anthropoid fossils date to the late Eocene and the early Oligocene.

d. Hominoids became widespread during the Miocene.

e. Hominids first appeared during the late Miocene or early Pliocene.

2. The Quaternary period has two epochs: Pleistocene and Holocene.


XII. Early Primates
A. Arboreal theory

1. Primates became primates by adapting to life in trees.

2. Enhanced sight (depth perception)

3. Grasping hands and feet


B. Visual predation hypothesis (Cartmill 1972, 1992)

1. Binocular vision, grasping hands and feet, and reduced claws developed because they facilitated the capture of insects.

2. Early primates first adapted to life in the bushy forest undergrowth and low tree branches.
C. Early Cenozoic Primates

1. The earliest primates date to the first part of the Cenozoic (65-54 m.y.a.).

2. The Eocene (54-38 m.y.a.) was the epoch of prosimians with at least 60 different genera in two families.

3. Anthropoids branched off from the prosimians during the Eocene.

a. Anthropoid eyes are rotated more forward compared to prosimians.

b. Anthropoids have a fully enclosed bony eye socket.

c. Anthropoids have a dry nose separate from the upper lip.

d. Anthropoids have molar cusps.


D. Oligocene Anthropoids

1. During the Oligocene (38-23 m.y.a.), anthropoids were the most numerous primates.

2. The parapithecid family may be ancestral to the New World monkeys.

3. The propliopithecid family may be ancestral to Old World monkeys, apes, and humans.


XIII. Miocene Hominoids
A. The earliest hominoid fossils date to the Miocene epoch (23-5 m.y.a.).
B. Proconsul

1. Proconsul was the most abundant anthropoid in the early Miocene.

2. Its teeth have similarities with modern apes, but below the neck the skeleton is more monkey-like.

3. Their teeth suggest that they ate fruits and leaves.

4. Proconsul probably contained the last common ancestor shared by Old World monkeys and the apes.

C. Later Miocene Apes

1. More than 20 species of primates have been discovered that date to the Late Miocene.

2. Middle and Late Miocene apes are often grouped into two families: Ramapithedidae and Dryopithecidae.

3. There are at least two ramapithecid genera: Sivapithecus and Gigantopithecus

a. Sivapithecus is now believed to be ancestral to the modern orangutan.

b. The orangutan line appears to have separated from the one leading to the African apes and humans more than 13 million years ago.

c. Gigantopithecus is the largest primate that ever lived, some standing over 10 feet tall and weighing 1,200 pounds.

d. Since it died out around 400,000 years ago, it coexisted with Homo erectus.


D. The Late Miocene

1. ???


2. Toumai is the name given to the 6 to 7 million year old primate skull found in northern Chad.

a. It may be the oldest human ancestor found yet.

b. The skull belonged to an adult male with a chimp-sized brain (320-380 cm3), heavy brow ridges, and a relatively flat, human-like face.

c. The find blends human and apelike features.

d. The find suggests that human ancestry was not confined to eastern and southern Africa.
XIV. Box: Saving the Orangutan
A. Dr. Birutá Galdikas has been working since the 1960s to study and preserve orangutans in Indonesia.
B. Orangutans are on the brink of extinction largely due to habitat loss.


LECTURE TOPICS



1. An illustration of homologies and analogies by using the various animals commonly familiar to undergraduates is most effective for demonstrating that difference. Such discussion also serves as the basis for explaining the importance of homologies in classification.
2. The variety of primate social groupings or organization should be categorized and presented in a simple tabular form for easy recall. The adaptive basis for the various forms should be made explicitly clear, particularly for the differences among the categories of mated couples, single male groups, groups in which relationships among female kin form the basis for social organization (female-bonded groups), and groups in which relationships among males form the basis for social organization (male-bonded groups; chimpanzees).
3. The adaptive roots of sexual dimorphism (competition among males for sexual access to females, in the case of most primates), both in external physical pressures and in social relations, should be explained carefully. How such adaptive pressures might or might not apply to humans should be considered.
4. There have been a number of studies done which investigate the nature of sex relations among nonhuman primates and attempt to draw some conclusions, for example, aboutthe origins of monogamy and marriage, sexual violence, and the origins of the sexual division of labor. Any of these issues is both interesting in its own right and useful as a case study illustrating the possibilities and limits for making generalizations across species. In particular, studies of male-to-female violence among nonhuman primates can be particularly useful for discussions on the “naturalness” of battering among humans.
5. Discuss the patterns of hominoid evolution during the Miocene.

SUGGESTED FILMS

Primates


1997 53 minutes

There are 234 primate species, and it seems that the more that is learned about them, the more humans come to understand themselves. Narrated by Armand Assante, this film explores the life of primates in their natural habitats. It discuss the social order, family life, diet, and mating habits of different primate species. A Discovery Channel Production.



Jane Goodall: A Life in the Wild


1990 31 minutes

This film presents the life and work of Dr. Jane Goodall. Through interviews with Dr. Goodall, we learn why she became a primatologist and how she came upon some of her more important findings. From Films for the Humanities and Sciences.



Washoe: Apes and Sign Language


1994 53 minutes

This film was made by the researchers who worked with Washoe and shows how Washoe acquired and used sign language. From Films for the Humanities and Sciences.



The Ape: So Human!


1998 41 minutes

This film presents the experiments by Allen and Beatrix Gardner, Sue Savage-Rumbaugh, and other primatologists related to chimpanzee behavior. Particular attention is paid to cognition, self-awareness, memory retention, language use, social behavior, and mating practices. From Films for the Humanities and Sciences.


Ancient Mysteries: Bigfoot

1994 50 minutes

Narrated by Leonard Nimoy, this film traces reports of Bigfoot sightings over thousands of years and discusses the current state of Bigfoot research. Presented by A&E Home Video.

Sasquatch Odyssey: The Hunt for Bigfoot


1999 60 minutes

This film presents a history of the search for Bigfoot, focusing primarily on four well-known Sasquatch researchers: Peter Byrne, John Green, Rene Dahinden, and Grover Krantz. Presented by Big Hairy Deal Films and Gryphon Productions Ltd., West Vancouver, BC, Canada.

MULTIPLE-CHOICE QUESTIONS
1. Which of the following primate traits are believed to have been selected for life in trees?
a. fewer offspring and bipedality

b. meat-eating and aggression

c. larger females and gentle males

d. fingernails (instead of claws) and soft fingertips

e. stereoscopic vision and an opposable thumb
2. Of the following primates, which are most likely to provide reasonable comparisons to humans?
a. lemurs

b. prosimians

c. New World monkeys

d. terrestrial primates

e. arboreal primates

3. Which of the following is not a hominoid?


a. gibbon

b. lemur


c. Homo sapiens

d. orangutan

e. siamang
4. Which of the following is shared by all anthropoids?
a. the ability to knuckle-walk and carry tools

b. bipedalism and one offspring born at a time

c. prehensile tails

d. a decrease in size of canines and an increase in size of molars

e. stereoscopic vision
5. Which of the following is used for putting organisms in the same taxon (zoological category)?
a. anthropometrics

b. homologies

c. only similarities that have evolved since the time of their common ancestor

d. analogies

e. all phenotypic similarities
6. Which of the following is an example of an analogy?
a. the mammary glands of dogs and the mammary glands of cats

b. similarities in chromosomal DNA between apes and humans

c. pentadactyly (five-digits on hands and feet) among baboons and macaques

d. dolphin fins and fish fins

e. bony eye sockets in chimps and similar structures in gorillas
7. What is the term for a trait that organisms have jointly inherited from a common ancestor?
a. analogy

b. homology

c. phenotype

d. allele

e. meiosis
8. Which of the following is not an adaptive trend in anthropoids?
a. stereoscopic vision

b. enhanced sense of touch

c. grasping hands and feet

d. decreased sociality

e. increased brain complexity
9. Ancient anthropoids began to have fewer offspring that required longer and more attentive care. What did this select for?
a. pair-bonding, which resembles the nuclear family, among 90% of present-day anthropoids

b. a greater capacity for brachiation

c. a greater reliance on nuts and tubers

d. increased reliance on arboreality (living in trees), which protected the young from predators

e. increased social complexity

10. Which of the following traits is not associated with primates?


a. reliance on smell as the main sense

b. stereoscopic vision

c. social groupings

d. grasping adaptations

e. brain complexity
11. Which of the following is not considered an anthropoid trend?
a. a shift from a moist muzzle and tactile hairs to fingers as the primary organs of touch

b. the evolution of a complicated visual system

c. an increase of cranial capacity relative to body size

d. an increase in hearing capacity at the expense of smell

e. a decrease in litter size
12. Recent research on chimpanzee eating habits indicates that
a. male chimps sometimes exchange meat for sex with females.

b. male chimps are exclusive herbivores.

c. chimpanzees have been observed “cooking” meat at volcanically heated springs.

d. while chimps do hunt a little, they get most of their meat by stealing it from predators.

e. chimpanzee hunting is the main reason New World monkeys are almost extinct.
13. lschial callosities and nonprehensile tails are characteristic traits of
a. Old World monkeys.

b. New World monkeys.

c. pongids.

d. prosimians.

e. tarsiers.
14. Which of the following primates is arboreal, active during the day, and has a prehensile tail?
a. prosimian

b. a New World monkey

c. an Old World monkey

d. a gibbon

e. extinct
15. Which of the following traits is not shared by humans and chimpanzees?
a. tool use

b. meat-eating

c. stereoscopic vision

d. high intelligence

e. estrus
16. Which of the following traits is not unique to gibbons and siamangs among the apes?
a. They are platyrrhines.

b. They are brachiators.

c. They exhibit little sexual dimorphism.

d. They live in primary groups consisting of one male, one female, and their preadolescent offspring.

e. They are completely arboreal.

17. New World monkeys are the only anthropoids who


a. have ischial callosities.

b. exhibit sexual dimorphism.

c. have prehensile tails.

d. are capable of brachiation.

e. have orthograde posture.

18. Which of the following traits is unique to humans?


a. social life

b. tool use

c. meat eating

d. food sharing

e. kinship
19. What is the greatest threat to endangered primates?
a. uniformitarianism

b. directional selection

c. intra-group predation

d. gene flow

e. loss of habitat
20. Which of the following traits is not a characteristic of anthropoids?
a. forward rotated eyes

b. bifrurcate merging lingual muscles

c. fully enclosed bony eye-socket

d. dry nose

e. four or five molar cusps
21. When do the first hominoid fossils appear?
a. during the Oligocene

b. during the Miocene

c. during the Pliocene

d. during the Pleistocene

e. during the Holocene
22. Why did the Old World monkeys thrive as the Miocene apes faded?
a. The Old World monkeys were superior at eating leaves.

b. The Miocene apes lacked opposable thumbs.

c. The Old World monkeys developed superior stone tools.

d. The Old World monkeys were bipedal.

e. The Miocene apes were too large for the environmental setting of the Miocene.
23. What species is believed to be ancestral to the modern orangutan?
a. Afropithecus

b. Kenyapithecus

c. Gigantopithecus

d. Sivapithecus

e. Dryopithecus

24. What species coexisted with Homo erectus in Asia


a. Afropithecus

b. Kenyapithecus

c. Gigantopithecus

d. Sivapithecus

e. Dryopithecus
25. Which of the following statements about Dryopithecus is not true?
a. It has been found in Europe and China.

b. It lived during the middle and late Miocene.

c. It had the Y-5 five cusp fissure pattern on its molars.

d. It subsisted on an exclusively carnivorous diet.

e. This taxon probably includes the common ancestor of the lesser apes and the great apes.

ESSAY QUESTIONS


1. What are the differences between human and nonhuman primate mating patterns?
2. Discuss trends that are characteristic of primate evolution. Indicate how these trends differentiate between the anthropoids and the prosimians.
3. Phylogenetically, who are our closest relatives? What evidence is used to support this relationship?
4. What do recent studies of primate social organization tell us about territorial behavior and aggression?
5. Based upon new evidence produced by studies of wild chimpanzees in various parts of Africa, what is the nature of chimpanzee hunting behavior? What are the implications of this behavior for such things as social organization and hypotheses about human origins?
6. What are the general trends in hominoid evolution during the Miocene? What derived hominoid traits appear during this time?

TRUE OR FALSE QUESTIONS


1. True or False Homologies are similarities between two species that have been jointly inherited from a common ancestor.
2. True or False Analogies are similarities that are shared by organisms that belong to the same genus.
3. True or False Opposable thumbs evolved as the early primates adapted to terrestrial life.
4. True or False Primates invest a lot of time and energy raising single offspring or small litters of offspring.
5. True or False Since primates are highly social animals, less care over a shorter period of time is provided to offspring.
6. True or False Platyrrhines are the New World monkeys.
7. True or False Sexual dimorphism tends to more pronounced in terrestrial species.

8. True or False Orangutans are found mostly in Madagascar.


9. True or False Chimpanzee social networks involve complex systems of dominance relationships.
10. True or False The spread of sickle-cell anemia to primate populations has drastically reduced the number of primate communities in the world.
11. True or False Learning, adaptive flexibility, and tool use are traits found only in the platyrrhines.
12. True or False Aggressive and violent behavior is not restricted to human populations.

13. True or False A key feature of primate evolution is the importance of smell over sight.


14. True or False The earliest primates were exclusive carnivores.
15. True or False Binocular vision improves depth perception.
16. True or False According to the arboreal theory, primates became primates by adapting to life in the savannah.
17. True or False The visual predation hypothesis argues that primates evolved elaborate ways of hiding themselves from their forest-based predators.
18. True or False It was during the Oligocene that anthropoids became the most numerous of the primates.
19. True or False Proconsul had ape-like teeth and an ape-like body.
20. True or False Sivapithecus is ancestral to modern orangutans.




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