Primates!

Specifically the EXTANT primates, i.e., the species that are still alive today: these include some prosimians, some monkeys, & some apes (-next: fossil hominins, who are extinct)

Taxonomy

What are primates?

Overview: What are primates?

•  Taxonomy of living things

•  Distinguishing primate characteristics

•  Primate taxonomy: distinguishing characteristics within the Order Primate…

•  Prosimians (Strepsirhines) –  Lorises –  Lemurs –  Tarsiers (?)

•  Anthropoids (Haplorhines) –  Platyrrhines

•  Cebids •  Atelines •  Callitrichids

–  Catarrhines •  Cercopithecoids

–  Cercopithecines –  Colobines

•  Hominoids –  Hylobatids –  Pongids –  Hominins

•  Subspecies •  Species •  Genus •  Family •  Infraorder •  Order •  Class •  Phylum •  Kingdom

Taxonomy: Hierarchical and Linnean (between Kingdoms and Species, but really not a totally accurate representation)

�Tree of life� -based on traits we think we observe -Beware anthropocentrism, the concept that humans may regard themselves as the central and most significant entities in the universe, or that they assess reality through an exclusively human perspective.

Taxonomy: Kingdoms (6 here)

Kingdom Animalia

•  Ingestive heterotrophs •  Lack cell wall •  Motile at at least some part of their lives •  Embryos have a blastula stage (a hollow ball of cells) •  Usually an internal digestive chamber

Phyla in Kingdom Animalia Phylum Meaning Group Acanthocephala Thorny head Thorny-headed worms

Acoelomorpha Without gut Acoels Annelida Little ring Segmented worms Arthropoda Jointed foot Arthropods Brachiopoda Arm foot Lamp shells Bryozoa Moss animals Moss animals, sea mats Chaetognatha Longhair jaw Arrow worms Chordata Cord Chordates

Cnidaria Stinging nettle Coelenterates Ctenophora Comb bearer Comb jellies Cycliophora Wheel carrying Symbion Echinodermata Spiny skin Sea Urchins Echiura Spine tail Spoon worms Entoprocta Inside anus Goblet worm Gastrotricha Hair stomach Meiofauna Gnathostomulida Jaw orifice Jaw worms Hemichordata Half cord Acorn worms Kinorhyncha Motion snout Mud dragons Loricifera Corset bearer Brush heads

Phylum Meaning Group

Mesozoa Middle animals Mesozoans

Micrognathozoa Tiny jaw animals —

Mollusca Thin shell Mollusks / molluscs

Myxozoa Slime animals

Nematoda Thread like Round worms

Nematomorpha Thread form Horsehair worms

Nemertea A sea nymph Ribbon worms

Onychophora Claw bearer Velvet worms

Orthonectida Straight swim

Phoronida Zeus' mistress Horseshoe worms

Placozoa Tubular animals

Platyhelminthes Flat worms Flat worms

Porifera Pore bearer Sponges

Priapulida Penis Priapulid worms

Rhombozoa Lozenge animal —

Rotifera Wheel bearer Rotifers

Sipuncula Small tube Peanut worms

Tardigrada Slow step Water bears

Xenoturbellida Strange flatworm —

Phylum Chordata

•  Hollow dorsal nerve cord •  Trends

–  Increasing cephalization –  Increased activity levels –  Increased predatory lifestyle

Phylum Chordata

•  Subphylum Urochordata (Tunicates: filter-feeding sea squirts)

•  Subphylum Cephalochordata (Lancelets) •  Subphylum Vertebrata (chordates with backbones)

–  Infraphylum Agnatha (jawless vertebrates) –  Infraphylum Gnathosomata (jawed vertebrates)

•  Superclass Osteichthytes (bony fishes) •  Superclass Tetraposa (four-legged vertebrates)

Subphylum Vertebrata •  BACKBONES •  Paired kidneys •  Heart, aorta •  Major transitions in some

ancestral vertebrates –  Superclass: Tetrapoda (4 limbs to

locomote on land) –  Amniotes

•  Shell, yolk, amnion, internal fertilization

•  Amnion = a membrane building the amniotic sac that surrounds and protects an embryo. It is developed in reptiles, birds, and mammals, but not in amphibians and and fish

Class mammalia •  Therapsids were transitional �mammal-like reptiles�

–  Occupied a nocturnal niche that dinosaurs didn’t dominate –  Increased metabolism to keep warm (bugs) –  Chewing (mastication); specialized teeth –  Legs under body to turn more easily

•  Mammals –  Survived age of dinosaurs –  First were tiny, nocturnal insectivores –  Escaped predation –  Survived global cooling

•  Adaptive Radiation(~ 0-65 mya) •  Three groups: •  Monotremes •  Marsupials •  Placental Eutherians

Mammals •  What features distinguish mammals? •  Hair made of keratin •  Active metabolism

•  endothermic •  hair and fat for insulation •  closed circulatory system + 4-chambered heart •  respiratory system •  connected to circulatory system •  diaphragm, a powerful muscle to enhance respiration

•  Reproductive characteristics •  Amniote egg without shell retained in uterus •  Viviparity •  Lactation

•  Cephalization •  Large Brains-key to human evolution •  Complex, social behaviors including parental care

Mammals •  Monotremes

•  Platypuses and Spiny Anteaters •  Lay reptilian, yolked eggs! •  But have hair and produce milk to nourish their

young •  Clear that modern monotremes are the survivors

of an early branching of the mammal tree; a later branching is thought to have led to the marsupial and placental groups

•  the egg is retained for some time within the mother, who actively provides the egg with nutrients. Monotremes also lactate, but have no defined nipples, excreting the milk from their mammary glands via openings in their skin

•  Extant in Australia and New Guinea, but widespread before that based on fossil data

•  Marsupials •  Opossums, kangaroos, koalas, sugar gliders •  Born early •  Complete embryonic development in pouch •  Nourished by milk (mammary glands) •  Diversified in Australia, few left in the Americas

(e.g., the opossum) •  Convergent Evolution with Placental (Eutherian)

Mammals

•  Eutherians (Placentals) •  Long pregnancy •  Complete development in utero •  Nourished by a placenta •  fetal membranes (chorion) and maternal tissues

intimately associated for gas exchange, nutrient supply, and waste removal!

More on Mammals

•  About 5400 species around today

•  Particular jaw feature defines mammals to paleontologists

•  Vivipary in many (but not limited to mammals—some sharks, etc.)

•  Most are placental •  Most are terrestrial

•  Order Macroscelidea: elephant shrews (Africa) •  Order Afrosoricida: tenrecs and golden moles (Africa) •  Order Tubulidentata: aardvark (Africa south of the Sahara) •  Order Hyracoidea: hyraxes or dassies (Africa, Arabia) •  Order Proboscidea: elephants (Africa, Southeast Asia) •  Order Sirenia: dugong and manatees (ctropical) •  Order Pilosa: sloths and anteaters (Neotropical) •  Order Cingulata: armadillos (Americas) (pink fairy armadillo here) •  Order Scandentia: treeshrews (Southeast Asia) •  Order Dermoptera: flying lemurs or colugos (Southeast Asia) •  Order Primates: lemurs, bushbabies, monkeys, apes •  Order Lagomorpha: pikas, rabbits, hares (Eurasia, Africa, Americas) •  Order Rodentia: rodents •  Order Erinaceomorpha: hedgehogs •  Order Soricomorpha: moles, shrews, solenodons •  Order Chiroptera: bats •  Order Cetartiodactyla: whales, dolphins and porpoises, even-toed

ungulates, including pigs, hippopotamus, camels, giraffe, deer, antelope, cattle, sheep, goats

•  Order Perissodactyla: odd-toed ungulates, including horses, donkeys, zebras, tapirs, and rhinoceroses

•  Order Pholidota: pangolins or scaly anteaters (Africa, South Asia) •  Order Carnivora: carnivores

•  Here: pink armadillo

Orders in the Class Mammalia

Order: Primates

•  What is a primate? •  Lots of diversity

Diversity

...200-300 species

Size

Lots of variation, but there are trends…

Primates •  Motor adaptations •  Large size variation across taxa •  Five digits on hands and feet: pentadactylism •  Mobile limbs •  Nails instead of claws (on at least one digit) •  Grasping digits with tactile pads (and fingerprints) •  Erect posture with extensive head rotation •  Rapid and precise muscle control •  Opposable hallus (big toe) and thumb (pincer grip) •  Prehensile hands and/or feet •  Well-developed clavicles (collarbones)

Primates •  Sensory adaptations •  Enlargement of eyes •  Color vision •  Binocular vision: overlapping fields of vision •  Stereoscopic vision and forward facing eyes:

–  neural wiring: sensory information from each eye relayed to both sides of brain

–  depth perception + accurate distance estimation

Primates •  Cranial Adaptations •  Reduced snout and olfaction •  Dental characteristics

–  Heterodont dentition (canines, incisors, molars and premolars)

–  Dental arcade (e.g., 2123 or 2133) –  In most groups, 32 or 36 adult teeth

Teeth

Primates •  Life history characteristics •  Delayed maturation •  Increased infant dependency •  Long gestation (pregnancy) •  Long lifespan •  Low reproductive rate •  Large, complex brain

Primates

•  Gregarious (many permanently live in groups) •  Male-female social associations •  Singleton births (in most species) •  Arboreal •  Diurnal •  Grooming (practical and social functions)

Primates •  How many species?

200-300(academic debates, hybrids…)

•  Future DNA sequencing may help sort it out, but maybe not, as genes may or may not address issues of human views on essentialism (c.f., Bloom�s book of recent)

•  New primate species are found every few years or so, in swampy areas that primatologists have a hard time getting around in

•  Extinction: pets and bushmeat (the next slide is rough)

Lavasoa Dwarf Lemur dicsovered in 2012 on Madagascar

Bushmeat and pets

Nonhuman (and undomesticated) animals are NOT meant to be pets

•  These are animals with social lives that are significant: primates tend to be social, are NOT domesticated, and do not make good �pets�

•  Food and �trophies� •  (…a Western construct?) •  Discussion?

Order Primates •  Prosimians (Strepsirhines)

–  Loris group –  Lemur group –  Tarsiers (?)

•  Anthropoids (Haplorhines) –  Platyrrhines

•  Cebids •  Atelines •  Callitrichids

–  Catarrhines •  Cercopithecoids

–  Cercopithecines –  Colobines

•  Hominoids –  Hylobatids –  Pongids –  Hominids

Order Primates •  Prosimians (Strepsirhines)

–  Loris group –  Lemur group –  Tarsiers (?)

•  Anthropoids (Haplorhines) –  Platyrrhines

•  Cebids •  Atelines •  Callitrichids

–  Catarrhines •  Cercopithecoids

–  Cercopithecines –  Colobines

•  Hominoids –  Hylobatids –  Pongids –  Hominids

Rhinarium = nose

wet, connected to upper lip strepsirhine

Dry, not connected to lip haplorhine

Primates •  The order Primates consists of two major suborders:

the Prosimians and the Anthropoidea. The prosimians were the first of the suborders to evolve; they are often called the "lower primates" The word prosimian literally means "pre-monkey."

•  Strepsirhines = prosimians •  Haplorhines = anthropoids (monkeys and apes)

Primates •  Prosimians •  Dental comb •  Postorbital opening •  Smaller brain •  Grooming claw •  Many nocturnal species •  More seasonal breeding

•  Anthropoids •  Dental comb absent •  Postorbital closure •  Larger brain •  Nails on all digits •  Few nocturnal species •  Less seasonal breeding

Postorbital Closure

Dental comb

Lemur catta

Prosimians •  Nose

–  Moist, naked rhinarium with the upper lip attached internally; greater reliance on olfaction than Haplorhine

•  Facial expression: less range possible (than for Haplorhini), partially because of attached upper lip; also far fewer structural featuress like tendons and muscles in their faces

•  Grooming "claw" on pedal digit for several (2nd digit) •  Teeth

–  dental formula (with some exceptions): 2133/2133 –  dental comb: elongated incisors used for grooming

•  Reproduction is generally seasonal •  Activity Period

–  often (but not always) nocturnal (or cathemeral) •  Many diurnal prosimians on Madagascar (where no diurnal monkeys or apes occur) •  nocturnal species have large eyes possessing tapetum lucidum

–  occasionally large & diurnal •  Infant care

–  nocturnal species often "park" their infants or cache them in nests while foraging –  cathemeral & diurnal species usually carry them

Prosimians

•  Lemuriformes: lemur group – Lemuridae (lemurs) –  Indriidae (indris, avahis, sifakas) – Daubentoniidae (aye-ayes)

•  Lorisiformes: loris group – Loridae (lorises, pottos, angwantibos) – Galagonidae (galagos)

•  Tarsiformes (tarsiers)???

Lemuriformes (the�lemur family�)

•  All only on Madagascar�s perimeter (extinct on mainland)

•  Lots of nocturnal species

Madagascar

• 22 or so mostly arboreal species • All members of lemur group are only on Madagascar�s perimeter (extinct on mainland)

Taxonomy: Prosimians (lemur group)

Family Body size* Activity Period General Social Pattern

"Special" Features

Cheirogaleidae Mouse & dwarf lemurs

Tiny Nocturnal Solitary - give birth to twins & cache them in nests - hibernate & accumulate fat seasonally (in tails)

Lepilemuridae Sportive lemurs

Small Nocturnal Solitary - coprophagy - sluggish

Lemuridae "True" lemurs

Medium Nocturnal Diurnal

Cathemeral

Solitary Social monogamy

Large multi-male, multi-female groups

- the ring-tailed lemur (Lemur catta) = only significantly terrestrial prosimian - female dominance over males sometimes

Indriidae Indri & sifaka & avahi

Small (avahi )

Medium (sifaka)

Large (indri )

Nocturnal (avahi)

Dirunal (indri, sifaka)

Solitary (avahi)

Multi-male, multi-female groups (sifaka)

Social monogamy (indri)

- female dominance over males sometimes

Lemur (lemur group)

Body size Activity Period

General Social Pattern

"Special" Features

Medium Nocturnal Diurnal

Cathemeral

Solitary Social

monogamy Large multi-male, multi-

female groups

- the ring-tailed lemur (Lemur catta) = only significantly terrestrial prosimian - female dominance over males sometimes

Lepilemur (�sportive� lemur; lemur group)

Body size Activity Period

General Social Pattern

"Special" Features

Small Nocturnal Solitary Coprophagy, sluggish

Mouse lemur (lemur group) Body size Activity Period General

Social Pattern

"Special" Features

Tiny Nocturnal Solitary - give birth to twins & cache them in nests - hibernate & accumulate fat seasonally (in tails)

Aye Aye (lemur group)

Body size Activity Period General Social Pattern

"Special" Features

Medium

Nocturnal Solitary - continuously growing incisors (rodent-like) - thin spindly middle finger = largest nocturnal primate

Others in the lemur group

Indri Avahi (�wooly lemur�)

Others in the lemur group

Sifaka (hoppers because of long, springy legs that let them jump 30 feet in trees) http://www.arkive.org/verreauxs-sifaka/propithecus-verreauxi/video-06b.html

Others in the lemur group Body size Activity Period General

Social Pattern

"Special" Features

Small (avahi )

Medium (sifaka)

Large (indri )

Nocturnal (avahi)

Dirunal (indri, sifaka)

Solitary (avahi)

Multi-male, multi-female groups

(sifaka)

Social monogamy (indri)

- female dominance over males sometimes

Lorisiformes (loris group)

Distribution (Africa and Asia)

Loris

Bushbaby (galago: loris-group prosimian)

Potto

Tarsier

Tarsier map

Tarsier issue in classification: prosimians or anthropoids?

order: Primates

suborder: Prosimii Anthropoidea

infraorder: Lemuriformes Lorisiformes Tarsiformes Tarsiformes ?

family: Lemuridae Indriidae Daubentoniidae Loridae Galagonidae Tarsiidae

species: lemurs indris avahis sifakas

aye-ayes lorises pottos

angwan-tibos

galagos tarsiers monkeys apes

humans

Anthropoids

•  Haplorhines •  Dry, unconnected noses •  Africa, Asia, South America •  Monkeys, apes and humans

–  ***prosimians are NOT �monkeys� •  Two groups

–  Catarhines –  Platyrhines

Rhine = nose

Catarrhine: Old World Primate Platyrrhine: New World Primate

Old/New World

Differences Between Platyrrhines and Catarrhines

Platyrrhine vs. Catarrhine Dentition: 2133 vs. 2123

(usually)

New World Monkeys

Ceboidea Family Subfamily Common Names Body size General Social

Pattern Some Special Features

Cebidae

Squirrel monkey

_______________

Capuchin monkey

Small

_________

Medium

Large, multi-male, multi-female groups

- strictly seasonal breeding

_______________________ - large brain

- tool use

-adaptable & resourceful (like Old World macaques)

- weakly prehensile tail

Callitrichidae

tamarins,

marmosets

small polyandry twins, high paternal investment,

reproductive suppression

Atelidae Pithecinae Uakaris & Sakis Medium

Socially monogamous

Other?

red-faced uakari:sexual selection? Swamp dwellers (hard to study)

Alouattinae Howler monkeys Large One-male, multi-female groups

("harems")

- LOUD howling

- prehensile tail (strong!)

Atelinae Spider monkey, wooly monkey, woolly spider

monkey

Very large Large fusion-fission communities

- prehensile tails (strong & dextrous!)

Interesting social patterns (kind of like chimps and bonobos)

Cebids vs. Callitrichids

Callitrichids

Pygmy marmoset (smallest primate)

juvenile golden lion tamarin

Callitrichids

Common Marmoset

Golden Lion

Tamarins

Emperor Tamarin

Cotton Top Tamarin

Moustached Tamarin

Atelines

Howler monkey

Red-faced uakari

Atelines: prehensile tails

Spider monkeys

Cebids

Squirrel monkey Capuchin monkey

Catarrhines

•  Cercopithecoids: the Old World monkeys – Cercopithecines – Colobinae

•  Hominoidea: the apes

Old World Monkeys

Cercopithecoidea

Cercopithecines

Hamadryas baboon

Gelada baboon

Japanese macaque

Cercopithecines: female-bonded

Celebes macaque

Colobines

Black and white colobus Snub-nosed langur

Colobines

Proboscis monkey

Colobines eat a lot of leaves

Hominoidea: the Apes Hylobatidae Hominidae

Category The “lesser” apes

The “great” apes The “human” apes

Common names

Gibbons and siamangs

Orangutans Gorillas Chimpanzees Bonobos humans

Distribution Southeast Asia

Borneo, Sumatra

-Lowland gorilla in West Central Africa -Mountain gorilla in volcanic mountains bordering Rwanda, Uganda, and Congo

Tropical rainforests and tropical forests of West, East, and North-central Africa

Central African Rainforests South of the big bend of the Congo River

Global (plus?)

Size Large (5-11 kg)

Huge (35-70 kg)

Huge (90-150 kg)

Huge (30-45 kg)

Huge (30-45 kg)

Huge

Grouping Pattern

Socially monogamous

Solitary Group (1 alpha male, his ‘harem,’ and their kids)

Large fission-fusion communities

Large fission-fusion communities

Large multimal/ multifemae communities

Ape Distribution

Brachiation gibbon

siamang

Duetting (territoriality)

Monogamy

Orangutan

Orangutan

Gorilla

Gorilla: mountain vs. lowland

Chimpanzee

Chimpanzee

Bonobo

Human

Primates •  Lots of morphological variation

–  Size, colors, dentition •  Lots of variation in social group structure (many males and

many females in a group vs. monogamous pairs, etc.) •  Lots of variance in social activity (solitary aye aye vs. the highly

gregarious capuchins and cercopithecines) •  Lots of variance in locomotion •  Lots of variance in diets •  Lots of variance in susceptibility to predation (e.g., large vs.

small animals)

•  What accounts for this variance?

Primate Behavioral Ecologists

•  Primatologists who try to figure out relationships between ecology, morphology, behavior, and sociality

•  Includes social variables (e.g., dominance and subordinance, fighting, mating, genetic relatedness), ecological variables (e.g., seasonal foods, the presence of predators), morphological variables (e.g., a very long gut), etc.

Some Examples

•  Colobines (OWM) and howler monkeys (NWM) eating leaves, but having very different energy levels

•  Male gorillas having proportionately larger teeth than females, even though they eat leaves, not meat

•  When newly joining a group, male langurs will selectively kill most or all infants who are still nursing, then immediately mate with the mothers (who agree to it!)

Goodall Films •  Four classic films from the

1960s of Goodall�s early work with Gombe (Tanzania—East Africa) chimpanzees

•  �Introduction to Chimpanzee Behavior�

•  �Infant Development� •  �Feeding and Food

Sharing� •  �Tool Using�

The Hadza •  The Hadza are an ethnic group in central Tanzania, living around Lake

Eyasi in the central Rift Valley and in the neighboring Serengeti Plateau. The Hadza number under 1000. Some 300-400 Hadza live as, hunter-gatherers much as they have for thousands, or even tens of thousands, of years; according to some anthropologists, they are of the last few functioning hunter-gatherers in Africa.

•  The Hadza are not closely related to any other people. While traditionally considered an East African branch of the Khoisan peoples, primarily because their language has clicks (see Pinker 167-168). The Hadza language appears to be an isolate, unrelated to or very distantly related to any other language. (e.g., 8 noun classes).

•  The Hadza subsist by hunting and gathering-foraging—which is what hominins (including our ancestors) did for millions of years during and prior to the evolution of anatomically modern humans

•  The lifestyle of the Hadza may be one of the world�s current best indicators of what life was like when humans evolved

•  Film (39 mins.) and how to take notes during a film

Hand Notes on �The Hadza�

Some notes on �The Hadza� •  Huts put up by women •  They are hunter-gatherers (HGs) •  Men hunt alone •  Women gather plant foods •  Hunt many impala •  Men gamble in downtime, winning and losing mainly arrows but also bees, honey; gambling losses do not cause significant

hardships ad arrows can be replaced by making new ones within hours to days •  No territoriality--people HG anywhere they want, unlike in most other HGs studied to the time the film was made •  All hunting by adult men is with bow and arrow; no traps, snares, or nets; the force require to pull a bow is strong--throws

off accuracy but implants arrow more deeply; 90% misses in hunting (often due to inaccuracy point above) •  Arrows with metal heads traded with other groups (for, e.g., honey) •  Majority of food from wild berries and roots/tubers, though they consider themselves meat-eaters •  Berries are abundant and many are eaten on the spot; food not stored past a day�s worth of eating (some seasonal

variation) •  Women about 2 hrs a day gathering all the food they need •  Baobob trees have tasty fruits (ground and cooked to a porridge); some trees better than others; berries are not like sweet

wildberries--harder to eat; roots and tubers cooked; water from underground sources and water trapped in tree hollows that are scarce during the dry season

•  Baobob and other locations have bees with honey and grubs •  Honey can be traded to other groups for western items like pots, tobacco, cloth, axes, iron/aluminum, beads •  One male duo did not find sufficient honey in one tree to walked a few miles to another known bee source •  Only collect enough food to last until the next day •  Dry months = more berries; wet months = more roots/tubers which are then more succulent •  Grandmas collect large tubers and share them with everyone in the camp--not just their specific families •  Hadza are nomads who only live in a particular camp for up to a few weeks •  Sometimes live on rock shelters--important returned-to shelters are on hills which provide a good vantage point of

surrounding areas •  An impala is killed; hidden from scavengers like vultures and hyenas until the hunter comes back with others to help carry

the carcass to camp •  The fat, not so much the meat, is what is prized in the impala--most hunted foods are high in protein but low in fat •  Boys practice hunting at young ages with wooden-tipped bows and arrows •  Boys catch a hyrax and use it to procure additional hyraxes, than kill & eat them all (no waste and yet no saving food)