P r i m a t e s

1
Primates
Nocturnal or Madagascar or both
S.E. Asia
S. America
Old World
2
Origins and Relations?
3
Evolution

• Primate adaptive themes exploitation of
  arboreal herbivory, arboreal locomotion, manual
  dexterity, stereoscopic vision, and complex
  social behavior and communication. (Vaughan et
  al., 2000)
• Increased manual dexterity (next 2 slides) may be
  correlated with increase in brain size.
• Mostly animals of the humid tropics, primates are
  an ancient (Cretaceous) Order, probably evolving
  in N. Am. (soon extinct) ?(Europe ?)Asia
  Africa? S.America
• It is useful to consider the Primates as 2
  Suborders
• The Strepsirhini (prosimian lemurs, bush
  babies, galagos, etc.).
• The Haplorhini (anthropoids monkeys, apes,
  people).
• The tarsier does not fit well with the
  anthropoids, but it is not a strepsirhine.

4
Prosimians (For each pair, hand is on left.)
5
Anthropoid primates (hands on left)
6
Taxonomic generalization (suborders)

• Strepsirhini (prosimians)
• Probably somewhat like earliest primates.
• Either Madagascar, or nocturnal, or both (all are
  tropical).
• Apparently do not compete effectively w/monkeys,
  etc.
• In contrast w/anthropoids
• More smell-oriented
• Less obviously social
• Seasonal breeders

• Haplorhini (anthropoids)
• Platyrrhini (New World)
• Catarrhini (Old World)
• Tarsiars are placed in this group largely because
  of molecular evidence.
• People are worldwide most other species are
  tropical.
• A commonly noted trend is toward behavioral
  (often social) adaptation rather than
  anatomical-physiological adaptation.

7
Ecological generalizations(regardless of
taxonomic position)

• Smaller primates
• For S/V reasons, must exploit high-energy foods.
• All eat arthropods plus occasional small
  vertebrates, which are difficult to collect in
  quantity. (Think protein.)
• Therefore most have other, reliable, sources of
  calories
• Fruit (very seasonal)
• Seeds (somewhat seasonal)
• Resins saps (less seasonal)

• Larger primates Most are either leaf specialists
  or fruit specialists
• Leaf eaters Energy-conserving second-rate
  fermenters that usually supplement calories with
  fruit. (These get plenty of protein if they make
  their calories.)
• Fruit eaters Plenty of energy must supplement
  protein with selected leaves or seeds or animal
  matter.

8
Tarsius Anthropoid life in the dark

• Formerly called a prosimian, now classed
  w/anthropoids, mostly on molecular evidence.
• The Family Tarsiidae includes 5 species in one
  genus.
• H-b of T. bancanus is 8.5-16cm mass is 100-150g.
• Nocturnal S.E. Asian primates eating only animal
  matter.
• Solitary/social ? an important concept

Reputedly has largest eyes (per body size) of any
animal even gives up eye-mobility for size. (Why
is size important?) Other night-vision
adaptations.
9
Uh, why do they call it a tarsiar?

• Diagram compares right feet of tarsier and brown
  lemur.
• Note extreme elongation of the calcaneum and
  navicular (tarsal bones) in the tarsier.
• The tarsier also has a radically different
  mechanism of foot-rotation so that it can grasp
  vertical stems with inverted foot.

10
Family Galagonidae (4g/11sp)

• All are restricted to Africa.
• All are nocturnal.
• All have long tails, large, mobile ears, and
  strong hind limbs (all are great leapers).
• All are highly vocal.
• Diets vary, but galagos eat lots of arthropods
  (from which they derive some calories all
  necessary protein). Most usually supplement this
  diet with fruit or with gums and resins.
• All are seasonal reproducers.

11
Galago sp. Lesser bush baby

• These African primates are slightly larger than
  tarsiars, and they have long tails.
• Primary habitat is woodlands.
• Mature more rapidly than other primates (XX
  sexually mature lt 1 yr).
• Except for mother with offspring, most galagos
  are solitary, but community is defined by scent
  marks.
• In some areas, gt 1 Galago species live
  sympatrically partitioning should be studied.

12
Otolemur crassicaudatus, greater bush baby

• Woodlands of southern Africa, including suburbs.
• Mass 1-1.5kg (XX slightly larger).
• gt 50 of calories from gums resins (especially
  in winter).
• Social structure poorly known
• XY said to be solitary XX said to move in
  groups?
• XX territories overlap, but not much XY
  territories encompass XX territories but dont
  overlap other XY territories?
• How do you resolve the contradictions?

13
Family Lorisidae (lorises potto)

• These are slow-moving nocturnal animals relying
  in part on gliding motion to avoid detection by
  predators.
• Unlike galagos, they do not jump.
• Wrists ankles specialized
• Great range of motion.
• Hands feet are zygodactylus.
• Special blood vessel storage channels in their
  hands and feet enable lorises to contract their
  muscles and hold tightly to a branch for hours,
  seemingly without fatigue or pain.
• Socially, these primates live in a world of
  smells.

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Loris tardigradus (slender loris)

• Range is s. 2/3 of India plus Sri Lanka.
• Mass is 200-450g.
• Diet is mostly arthropods.
• Solitary but social (!)
• Foraging usually solo, sleeping often not
• XY mark territories by fecal sign posts
• XX leave urine trails w/social information.

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Nycticebus coucang, slow loris

• Deep-forest animal of S.E. Asia.
• H-b 25-35cm, mass 1.5kg.
• Diet 50 fruit w/ protein hits from small
  animals.
• Social structure ecological relationships to
  other loris sps are poorly known in wild.

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Perodicticus potto, the potto

• Nocturnal animal of W. African rainforest canopy.
• H-b 25-35cm mass 1kg.
• Basis of diet is fruits, plus gums and small
  animals for protein hits.
• Young learn to eat by foraging with mother.
• Defensive behavior.

17
The Lemuriform primates

• All of these are restricted to Madagascar (see
  next slide).
• Here are some Family sketches
• Cheirogaleidae 4 genera of nocturnal primates
  ecologically more similar to African prosimians
  than to other Madagascar lemurs.
• Megaladapidae Has only 1 living species, a
  leaf-fermenter, but formerly included huge
  terrestrial types filling artiodactyl niches.
• Lemuridae 4g/10s. Most species are diurnal.
  Often partition habitat finely mostly
  monkey-convergents.
• Indridae 3g/5s. Large leaf-eaters, mostly
  diurnal. Some extinct members were large,
  terrestrial browsers.
• Daubentoniidae The aye-aye is a strange
  nocturnal insect-eater, a woodpecker-convergent.

18
Alternative Evolution Madagascar invaded by only
6 mammal Orders Insectivores, carnivores,
rodents, bats, artiodactyls primates.
19
Patterns of Malagasy Extinctions
20
Aye-ayeDaubentoniidae

• Originally classified as a rodent, this nocturnal
  primate occupies woodpecker niche on Madagascar.
• Note chisel teeth, long middle finger, and use of
  the finger in feeding.
• Like much other Malagasy fauna, aye-ayes are
  endangered by habitat destruction.

21
Brown ringtail lemurs (Lemuridae 5 genera,
about 25 species)

• Eulemur fulvus
• Mostly arboreal.
• Leaves fruit.
• Flexible, multi-XY groups that can fracture into
  subgroups.
• Lemur catta
• Mostly terrestrial.
• Dietary generalists.
• Group structure is organized around XX residents
  which are entirely dominant over transient XY.

22
Behavioral adaptability in the Lemuridae
23
Fat-tail mouse lemurs (Cheirogaleidae 4
genera 17 species)

• Cheirogaleus medius (left)
• 200g (medium rat).
• Eat fruit protein hits.
• Only hibernating primate.
• Microcebus murinus (right)
• 30g (medium mouse)
• Rainforest-edge species
• Foraging (for anything) mostly solitary, but XX
  day-nest together.
• Heavy depredation pressures rapid maturity
  twice-annual litters of 2-4.

24
Anthropoid primates Phylogeny general trends

• Increased body-mass
• Increased brain-mass/body-mass
• Longer life
• Increased manual dexterity
• Specialization of eyes
• Binocular vision (at expense of olfaction)
• Increased density of cones
• Emphasis of visual acuity over light-gathering

25
Notes on sociality in higher primates

• Primatologists are abundant and specialized
  often they use a vocabulary of their own, naming
  many types of social structures.
• Most general mammalogists agree that
• Primate groups vary among and within species.
• Most social organizations seen in primates have
  also been documented in other mammal species.
• Intraspecific variation is often correlated
  w/habitat.
• Territorial XY primates appear more attached to
  XXs than to land.
• XX among primates are more likely (than XX among
  other mammals) to leave natal groups (but
  remember equids?).
• Human social organization varies is almost
  unique.

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New World Monkeys

• Ancestor rafted from Africa.
• Groupings
• MT small, fruit exudate eaters.
• CS Highly adaptable fruit eaters.
• NT Unripe fruit, leaves, insects.
• US Seed eaters.
• H Leaf eaters.
• SW Large, brachiating fruit eaters.

27
Callitrichidae (marmosets our first anthropoid
Family)

• 4 genera/26species of small monkeys confined
  almost entirely to South American forests.
• Evolutionary origins controversial
• Small size is primitive condition?
• Small size is specialization for insect resin
  diet?
• Little sexual dimorphism.
• Group nucleus is one breeding pair (often
  monogamous) that is often assisted by
  stay-at-home older kids.
• Litter size is typically 2 (dizygotic twins
  shared placenta) offspring receive extensive
  family care.

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Marmosets tamarins (Callitrichidae)

• Golden-chested marmoset
• Mass lt 500g squirrel-like in activity.
• Most calories from resins.
• Sociality reproduction.
• Golden lion tamarin
• Mass c. 500-750g.
• Most calories from fruits.
• Sociality reproduction.

29
New World Monkeys Cebidae

• 11 genera, 58 species, small to medium size.
• Vast majority South American w/ some species
  extending into Central America and a few crossing
  into North America (in s. Mexico).
• Evolutionary origins unclear, but emerging
  consensus holds with rafting from Africa?
• All species but one are diurnal.
• Tail typically long, often prehensile.
• Major ecological differences are between
  fruit-eaters and leaf-eaters.

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Squirrel monkeys (Saimiri)

• Not much larger than marmosets (0.7-1.2kg).
• Social and arboreal omnivores.
• Large groups, poorly studied
• XX maturity _at_ 2 yrs sisterhood, aunting.
• XY Loose dominance hierarchy mediated by
  physiological changes.

31
Cebus monkeys

• Cebus has the largest brain/body ratio of any New
  World primate.
• Most diverse diet of any New World primate.
• One of few mammals in which XXs often outlive
  repro. years.
• Members of this genus are highly adaptable
• In the wild home range, occasional tool-use,
  social organization.
• In captivity, general train-ability.

C. albiniforns (left) C.
capucinus (right)
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Dusky Titi Monkey (Callicebus moloch)

• Diet is mostly leaves and green fruit,
  supplemented by ripe fruit and insects.
• Titis are basically monogamous, bearing young
  early in rainy season.
• Although they do well in zoos, these monkeys are
  not well known in the wild.

33
Night Monkey (Aotus trivirgatus)

• Sometimes called owl monkeys, these are the most
  nocturnal anthropoid primates.
• Diet is fruit, leaves, insects ( small verts).
• Basically monogamous.
• The taxonomy of this monkey is confusing some
  authorities name up to 9 species.
• This species is of considerable medical interest
  because owl monkeys are immune to malaria.

34
Uakaris (Cacajao calvus)

• These are poorly known monkeys with short tails,
  shaggy bodies, naked heads.
• Mass is 3-4kg w/XY slightly heavier than XX.
• Habitat makes study of uakaris almost impossible
• Multi-XX, multi-XY groups w/XY dominance.
• Food may be primarily seeds.

35
Sakis (Pithecia sps.)

• Sakis are medium-small monkeys (1.5-2kg).
• Diet includes fruit but is probably dominated by
  seeds and nuts (see teeth).
• Social structure is not well known but involves
  breeding pairs. (Sakis are sexually dichromatic.)
• These are great leapers in mature forests.

36
Howlers (Alouatta sps.)

• The most common monkeys in most Neotropical
  systems.
• XX mass 4-5kg XY 6.5kg.
• Cautious, lazy leaf-eaters marking territory by
  vocalizations.
• Group size reflects resource density.
• Groups are 1-XY harems within which XX rankings
  are age-inverted.

37
Wooly Spider Monkeys

• These are the largest New World monkeys h-b
  length to 60cm mass to 12kg.
• Diet is mainly fruit, supplemented for protein.
• Group structure is highly flexible, varying with
  habitat and season.
• Sexual dimorphism is minimal though males do
  have dominance hierarchies, they are loose
  generally age-structured.

38
Old World Primates
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Family Cercopithecidae

• The most successful primate Family (18g/81s)
• Africa East, South, Southeast Asia. Extending
  beyond tropics in southern Africa, China, Tibet,
  and Japan.
• Mass ranges from 1.5 to gt 50kg.
• Most are diurnal all are highly social.
• Ecologically divisible into 2 groups
• Colubinae, the Old World leaf eaters
• Cercopithecinae, the Old World generalists.

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African leaf-eaters (e.g., Colobus guereza)

• Mass XX 8kg XY 12kg.
• Central Africa habitat is rainforest, gallery
  forest, well-structured secondary forest.
• The most capable fermenters among African
  primates (all primates?).
• Multi-XY groups (8-15) w/dominance hierarchies.
• Small (c. 18ha) ranges defended against other
  groups.

41
Asian leaf-eaters (e.g., Semnopithecus entellus)

• Hanuman langurs are south-Asian.
• XX 12kg XY 18kg.
• Diet 35 leaves, 25 fruit 40 misc.
• Social structure changes as function of monkey/
  resource density
• Low multi-XY groups.
• High 1-XY harems plus bachelor groups that take
  over harems (lion pattern).

42
Cercopithecus Chlorocebus aethiops

• Found in southern African open woodlands.
• XX 3kg XY 4kg.
• Group size ecologically flexible (usually 15-25)
  with multiple XY and dominance hierarchies.
• XX stay in natal group XY out-migrate.
• CIVs
• Each species of Cercopithecus (sensu lato) may
  have its own.
• About 50 are non-symptomatic carriers.

43
Baboons (Papio hamadryas)

• Baboons are large, pan-African monkeys living in
  large, intricately-structured social groups
  evolved in part for collective defense.
• Sexual dimorphism is extreme XX 15kg XY to
  50kg.
• Diet includes anything that provides good
  nutrition.

44
Further baboon notes

• Group structures and subspecies/habitat
  differences
• Effectiveness of group defense where needed.
• Southern African savannas 20-50 animals group
  defense in a predator-rich environment.
• Ethiopia Few big, safe sleeping areas ( thus
  groups of several hundred baboons) sparse
  resources ( thus foraging groups of 1-2XY XX),
  residual typical groups.
• Life within a group
• XY hierarchy obvious important XX hierarchy
  subtle and more important for eventual status of
  offspring. Until mature, (c. 4 years) XY remain
  subordinate to XX.
• Godfather relationships rare but potentially
  important.
• Political behavior, troop shake-ups, and XX
  behavior.

45
Other baboon-like monkeys

• Mandril (Mandrilus spinx) This is a large
  monkey with brightly-colored face hindquarters,
  poorly known in the wild, that fills a
  baboon-type niche in African rainforests. Groups
  are smaller than among baboons.
• Gelada (Thercopithecus gelada) This is a big
  monkey of the Amhara Plateau. Sociality looks at
  first like that of baboons but XX are socially
  more important. Perhaps this reflects the fact
  that gelada XX stay with their natal groups while
  baboon XX out-migrate.

46
Macaques (Asian generalists)

• Macaques include one genus (Macaca) 16 sps.
• Diets, social structures, and habits vary by
  habitat, but all are adaptable. All include
  multi-XY groups w/ XY XX dominance hierarchies.
• Long-term studies on Japanese macaques indicate
  group innovations are brought in by young XXs.
• Example is crab-eating macaque, Macaca
  fascicularis. Group size is 10-50 food mostly
  fruit.

47
Macaca nemestrina sociality, 1

• Adult XX with offspring.
• As with baboons, macaque babies inherit to some
  degree the status of their mothers.
• Also, the status of a XX with infant is
  temporarily raised.
• And XXs who are consistently successful mothers
  often have higher status.

48
Macaca nemestrina sociality, 2

• Adult XY, pucker-threat.
• Note the crouch, the extended lips, and the
  forward-directed posture, which are diagnostic
  for several types of threat.
• The pucker-threat is an assertion of dominance
  that typically elicits either submission or a
  reciprocal challenge (and then possibly a fight).

49
Macaca nemestrina sociality, 3

• Adult XX, pucker-threat.
• XX almost always respond submissively to XY
  threats, but macaques are also individuals.
• This XX (youve seen her before) is able to
  reciprocate the XY threat because (1) hes
  low-ranking, (2) shes usually high-ranking, and
  (3) she has an infant with her.

50
Macaca nemestrina sociality, 4

• Male-male mount.
• In primate societies things that appear to be
  about sex are often about power.
• To appease a higher-ranking XY, a lower-ranking
  XY acts submissively, presenting sexually (an
  XX gesture).
• The higher-ranking XY accepts, with a symbolic
  mount.
• Note the young animal looking on and becoming
  socialized.

51
Macaca nemestrina sociality, 5

• Low-status adult XY, fear-yawn.
• The lowered eyes, raised brows, retracted head
  indicate fear, as does the yawn itself.
• The exposed canines indicate that attack would
  nevertheless risk injury.
• Note that some facial gestures have been
  conserved for MY in primates.

52
Macaca nemestrina sociality, 6

• Coalition of brothers.
• As with other social animals (e.g., lions),
  displacement of group leaders is often
  accomplished by coalitions.
• In macaques, coalitions almost always involve
  (half-) brothers because most lasting social
  connections are established through mothers.

53
The Ape Grade

• A general evolutionary trend among anthropoid
  primates is increasing reliance on vision at the
  expense of smell. The trend culminates in apes
  and is associated w/increased hand-eye
  coordination enlargement of the brain.
• Ape-grade primates lack external tails. In all
  except people, arms are longer than legs.
• Apes have long inter-birth intervals extended
  parental care (both anthropoid trends).
• All ape-grade primates (except people) are
  pan-tropical and endangered by direct
  exploitation and/or habitat destruction.

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Family Hylobatidae

• This Family includes gibbons siamangs (1 genus,
  11 species) picture example is Symphalagus
  syndactylus.
• These are small (10-12kg), tailless, fruit-eating
  apes of S.E. Asian rainforests.
• Monogamous fidelity is extreme. Inter-birth
  intervals are c. 3 years. 2-parent care is
  extensive (6-7 years).
• First birth is at about 9 years.
• Pairs mark territory by brachiating displays
  duets.

55
Orangutans (2 similar species)

• Pongo pygmaeus, from Kalimantan, is illustrated
  another sp. on Sumatra.
• XX ave. 40kg wild XY twice as heavy.
• Adult XY avoid other XY use roars to maintain
  territories overlapping ranges of gt 1 XX.
• XX are at first repro is about 12-15yrs. Only
  cohesive social unit is mother offspring.

56
Common chimp (Pan troglodytes)

• Chimps are forest animals of Central West
  Africa.
• XX 35kg XY 45kg.
• Diet varies but is usually dominated by fruit.
  (Hunting traditions in some groups.)
• XX sexually mature _at_ 14 inter-birth interval c.
  5 years.
• Unique because XY-XY bonds form backbone of
  social groups (XX out-migrate).
• Chimps share c. 98 of genes with people.

57
Bonobo (Pan paniscus)

• The bonobo, or pygmy chimp, inhabits central
  rainforest along the Congo basin.
• Anatomically and genetically similar to common
  chimp
• but radically different socially
• Sex seems to be the glue that holds bonobo
  society together.
• XXs are migratory gender, but relationships
  among them are strong.
• Even adult XYs appear to take status partly from
  mothers.

58
Gorillas (Gorilla gorilla 3ssp.)

• Gorillas are, on ave., the largest primates XX
  85-95kg XY c. 165kg.
• Habitat is dense forests, swamps, etc.
• 2 lowland ssps eat fruit mountain ssp eats
  leaves shoots.
• Group size typically 5-20 most include 1 (2)
  large XY plus XXs young.
• Inter-birth interval 3-5 years.
• XX XY out-migrate.

59
People (Homo sapiens)

• The most abundant primate species.
• Large African primate most closely related to
  chimps but radically divergent in anatomy
  behavior.
• People are world-building (as opposed to
  world-accepting) animals relying on artificially
  constructed ecosystems for almost all material
  needs.