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Goodall Films

• Four classic films from the 1960s of Goodalls early work with Gombe (Tanzania —East Africa) chimpanzees • Introduction to Chimpanzee Behavior • Infant Development • Feeding and Food Sharing • Tool Using !

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

Diversity

...200$300&species&

What are primates? Overview: What are primates?

• Taxonomy of living • Prosimians (Strepsirhines) – things – • Distinguishing – (?) • Anthropoids (Haplorhines) – Platyrrhines characteristics • Cebids • Atelines • Primate taxonomy: • Callitrichids distinguishing characteristics – Catarrhines within the Primate… • Cercopithecoids – Cercopithecines – Colobines • Hominoids – Hylobatids – Pongids – Hominins Taxonomy: Hierarchical and Linnean (between Kingdoms and Species, but really not a totally accurate representation)

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

Tree of life -based on traits we think we observe -Beware anthropocentrism, the concept that may regard themselves as the central and most significant entities in the universe, or that they assess reality through an exclusively 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 Phylum Meaning Group Acanthocephala Thorny head Thorny-headed worms Mesozoa Middle Mesozoans Acoelomorpha Without gut Acoels Micrognathozoa Tiny jaw animals —

Annelida Little ring Segmented worms Thin shell Mollusks / molluscs

Arthropoda Jointed foot Myxozoa Slime animals Brachiopoda Arm foot Lamp shells Nematoda Thread like Round worms Bryozoa Moss animals Moss animals, sea mats Nematomorpha Thread form Horsehair worms Chaetognatha Longhair jaw Arrow worms Nemertea A sea nymph Ribbon worms Chordata Cord Onychophora bearer Velvet worms Cnidaria Stinging nettle Coelenterates Orthonectida Straight swim Ctenophora Comb bearer Comb jellies Phoronida Zeus' mistress Horseshoe worms Cycliophora Wheel carrying Symbion Placozoa Tubular animals Echinodermata Spiny skin Sea Urchins Platyhelminthes Flat worms Flat worms Echiura Spine tail Spoon worms Porifera Pore bearer Sponges Entoprocta Inside anus Goblet worm Priapulida Penis Priapulid worms Gastrotricha Hair stomach Meiofauna Gnathostomulida Jaw orifice Jaw worms Rhombozoa Lozenge — Hemichordata Half cord Acorn worms Rotifera Wheel bearer Rotifers Kinorhyncha Motion snout Mud dragons Sipuncula Small tube Peanut worms Loricifera Corset bearer Brush heads 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 (animals with backbones) – Infraphylum Agnatha (jawless ) – 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 , but not in amphibians and and fish Class mammalia

• From synapsids…therapsids • Therapsids were transitional -like reptiles – Nocturnal niche that dinosaurs didnt dominate – Increased metabolism to keep warm (bugs) – Chewing (mastication); specialized teeth – Legs under body to turn more easily – Oligokyphus (but some cranial features make it not a mammal) • 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 • Monotremes • Platypuses and Spiny Anteaters • Lay reptilian, yolked eggs! Mammals • 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, , koalas, sugar gliders • Born early • Complete embryonic development in pouch • Nourished by milk (mammary glands) • Diversified in Australia, few left in the (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 Orders in the Class Mammalia

• 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 Primates

• Mivart, 1873: "Unguiculate, claviculate placental mammals, with orbits encircled by bone; three kinds of teeth, at least at one time of life; brain always with a posterior lobe and calcarine fissure; the innermost digit of at least one pair of extremeties opposable; hallux with a flat nail or none; a well-developed caecum; penis pendulous, testes scrotal; always two pectoral mammae." 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 (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 (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 (canines, incisors, molars and premolars) – 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., Blooms book of recent) • New primate species are found every few years or so, in swampy areas that primatologists have to slough through…this work is not easy and cute, as we might think when we think about cute monkeys • : and bushmeat (the next slide is rough) 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) – group – group – Tarsiers (?) • Anthropoids (Haplorhines) – Platyrrhines: New World Monkeys • Cebids • Atelines • Callitrichids – Catarrhines: Old World Monkeys and Apes • 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 =

wet, connected to upper lip Dry, not connected to lip strepsirhine 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 literally means "pre-." • Strepsirhines = prosimians • Haplorhines = anthropoids (monkeys and apes) Primates

• Prosimians • Anthropoids • Dental comb • Dental comb absent • Postorbital opening • Postorbital closure • Smaller brain • Larger brain • • Nails on all digits • Many nocturnal species • Few nocturnal species • More seasonal breeding • 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 ), 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 – 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

: lemur group – (lemurs) – (indris, avahis, sifakas) – Daubentoniidae (aye-ayes) • Lorisiformes: loris group – Loridae (lorises, pottos, angwantibos) – Galagonidae () • Tarsiformes (tarsiers)??? Lemuriformes (the lemur family) • All only on Madagascars perimeter (extinct on mainland) • Lots of nocturnal species Madagascar

• 22 or so mostly arboreal species • All members of lemur group are only on Madagascars perimeter (extinct on mainland) Taxonomy: Prosimians (lemur group)

Family Body size* Activity Period General "Special" Features Social Pattern

Cheirogaleidae Tiny Nocturnal Solitary - give birth to twins & cache Mouse & dwarf lemurs them in nests - hibernate & accumulate fat seasonally (in tails) Lepilemuridae Small Nocturnal Solitary Sportive lemurs - coprophagy

- sluggish Lemuridae Medium Nocturnal Solitary - the ring-tailed lemur (Lemur "True" lemurs Diurnal Social monogamy catta) = only significantly Cathemeral Large multi-male, multi- terrestrial prosimian female groups - female dominance over males sometimes Indriidae Small Nocturnal Solitary - female dominance over Indri & sifaka & avahi (avahi ) (avahi) (avahi) males sometimes

Medium Dirunal Multi-male, multi-female (sifaka) (indri, sifaka) groups (sifaka) Large (indri ) Social monogamy (indri) Lemur (lemur group)

Body size Activity General "Special" Period Social Features Pattern

Medium Nocturnal Solitary - the ring-tailed Diurnal Social lemur (Lemur Cathemeral monogamy catta) = only Large multi- significantly male, multi- terrestrial female groups prosimian - female dominance over males sometimes Lepilemur (sportive lemur; lemur group)

Body size Activity General "Special" Period Social Features Pattern Small Nocturnal Solitary Coprophagy, sluggish Mouse lemur (lemur group)

Body size Activity Period General "Special" Social Features Pattern Tiny Nocturnal Solitary - give birth to twins & cache them in nests - hibernate & accumulate fat seasonally (in tails) Body size Activity Period General "Special" Social Features Pattern

Medium Nocturnal Solitary - continuously Aye Aye growing incisors (rodent-like) - thin spindly (lemur group) 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 "Special" Social Features Pattern Small Nocturnal Solitary - female (avahi ) (avahi) (avahi) dominance over males sometimes Medium Dirunal Multi-male, multi- (sifaka) (indri, sifaka) female groups (sifaka) Large (indri ) Social monogamy (indri) Lorisiformes (loris group)

Distribution (Africa and Asia) Loris Bushbaby (: loris-group prosimian) Potto 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 aye-ayes lorises galagos tarsiers monkeys avahis pottos apes sifakas angwan- humans tibos

Anthropoids

• Haplorhines • Dry, unconnected • Africa, Asia, • 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: 2132 and 2133 vs. 2123 (usually) New World Monkeys Ceboidea Family Subfamily Common Names Body size General Social Some Special Features Pattern Small Large, multi-male, multi-female groups - strictly seasonal breeding ______

- large brain ______Capuchin monkey Medium - tool use -adaptable & resourceful (like Old World macaques)

- weakly small polyandry tamarins, twins, high paternal investment, reproductive suppression marmosets Pithecinae Uakaris & Sakis Medium Socially monogamous red-faced uakari:sexual Other? selection? Swamp dwellers (hard to study)

Alouattinae Howler monkeys Large One-male, multi- female groups - LOUD howling ("harems") - prehensile tail (strong!)

Atelinae , Very large Large fusion-fission - prehensile tails (strong & wooly monkey, communities dextrous!) woolly spider Interesting social patterns (kind monkey of like chimps and bonobos) Cebids vs. Callitrichids

cebids Callitrichids gross characterization “true” monkeys “squirrel like” monkeys body size medium tiny dental formula 2133 (top and bottom) 2132 (top and bottom) nails and claws nails on all digits claws on all digits except hallux (big toe) births Singletons twins male care of infants Rare and indirect present and direct in most species alloparental care Absent except in Perry’s present in most species white faced capuchins

Callitrichids

juvenile golden lion tamarin

Pygmy marmoset (smallest primate) Callitrichids Golden Lion Tamarins

Emperor Tamarin Moustached Tamarin

Cotton Top Tamarin Common Marmoset Atelines

Red-faced uakari

Howler monkey Atelines: prehensile tails

Spider monkeys Cebids

Squirrel monkey Capuchin monkey Catarrhines

• Cercopithecoids: the Old World monkeys – Cercopithecines – Colobinae • Hominoidea: the apes Old World Monkeys Cercopithecoids Cercopithecines

Gelada baboon

Hamadryas 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

Category The “lesser” The “great” apes The apes “human” apes Common and Orangutans Gorillas Chimpanzees Bonobos humans names siamangs

Distribution Southeast Borneo, -Lowland gorilla in Tropical Central African Global Asia Sumatra West Central Africa rainforests and Rainforests (plus?) -Mountain gorilla in tropical forests of South of the big volcanic mountains West, East, and bend of the bordering Rwanda, North-central Congo River Uganda, and Congo Africa Size Large Huge Huge Huge Huge Huge (5-11 kg) (35-70 kg) (90-150 kg) (30-45 kg) (30-45 kg)

Grouping Socially Solitary Group (1 alpha male, Large fission- Large fission- Large Pattern monogamous his ‘harem,’ and their fusion fusion multimal/ kids) communities communities multifemae communitie s Distribution Brachiation

siamang Duetting (territoriality) Monogamy Orangutan Orangutan Gorilla Gorilla: mountain vs. lowland Chimpanzee Chimpanzee Bonobo Human