Primates! Specifically the EXTANT primates, i.e., the ones who 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 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 humans may regard themselves as the central and most significant entities in the universe, or that they assess reality through an exclusively human perspective. Dendrogram (cladistics based on genes) Taxonomy: Kingdoms (5 here) 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 animals Mesozoans Acoelomorpha Without gut Acoels Micrognathozoa Tiny jaw animals — Annelida Little ring Segmented worms Mollusca Thin shell Mollusks / molluscs Arthropoda Jointed foot Arthropods 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 Chordates Onychophora Claw bearer Velvet worms Cnidaria Stinging nettle Coelenterates Orthonectida Straight swim Ctenophora Comb bearer Comb jellies Cycliophora Wheel carrying Symbion Phoronida Zeus' mistress Echinodermata Spiny skin Sea Urchins Placozoa Tubular animals Echiura Spine tail Spoon worms Platyhelminthes Flat worms Flat worms Entoprocta Inside anus Goblet worm Porifera Pore bearer Sponges Gastrotricha Hair stomach Meiofauna Priapulida Penis Priapulid worms Gnathostomulida Jaw orifice Jaw worms Rhombozoa Lozenge animal — 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 — Horseshoe worms 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 • • • • • • • • From synapsids…therapsids Therapsids were transitional “mammal-like reptiles” – Nocturnal niche that dinosaurs didn’t 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 • • • • • • • Marsupials • • • • • • • 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 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! Mammals Mammals • • • • • • • • • Milk/mammary glands Hair Fur Specialized teeth Neocortex Warm-blooded, circulation, heart 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 Class Mammalia • • Subclass Prototheria (monotremes: platypuses and echidnas: lay eggs but have hair and produce milk) Subclass Theria - live-bearing mammals – Infraclass Metatheria - marsupials – Infraclass Eutheria - placentals • • • • • • • • • • • • • • • • • 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) 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 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 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) – 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 to slough through…this work is not easy and cute, as we might think when we think about “cute monkeys” • Extinction: pets 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? Suborder Infraorder Superfamily Family Subfamily Lemuridae (true lemurs) Lemuriformes Indriidae Daubentoniidae Prosimii (prosimians) Loridae Lorisiformes Galagonidae Tarsiformes Tarsiidae Callithricidae Platyrrhini (New World monkeys) Ceboidea Cebidae Atelidae Anthropoidea (anthropoids) Catarrhini (Old World monkeys, apes and humans) Cercopithecoi dea (Old World monkeys) Cercopithecin ae Cercopithecida e Colobinae Hylobatidae Hominoidea (apes and humans) Pongidae Hominidae Common Names Distribution ring-tailed, ruffed, and mouse lemurs, etc. Madagascar and Comoro Islands indris, avahis, and sifakas Madagascar aye-ayes Madagascar lorises, pottos, and angwantibos lorises--India and Southeast Asia; others-Africa galagos (or bush babies) sub-Saharan Africa and Zanzibar tarsiers Philippines, Borneo, Celebes Islands, and Sumatra marmosets and tamarins Panama and north and eastern South America squirrel and owl monkeys, capuchins, etc. Central America and north and eastern South America howler and spider monkeys, etc. Central America and north and eastern South America guenons, vervets, baboons, macaques, etc. guenons and baboons-Africa; macaques-northwest Africa, Gibraltar, South and East Asia colobuses, langurs, and proboscis monkeys colobuses-- Central Africa; langurs-- India and Southeast Asia; proboscis monkeys-Borneo gibbons and siamangs Southeast Asia orangutans Sumatra and Borneo chimpanzees, bonobos, and gorillas Central and West Africa humans throughout the world Taxonomy Taxonomy: How do we classify animals? A. Example of classifying the dusky titi monkey, Callicebus moloch: (suffixes noted in parentheses) Order: Primates Suborder: Haplorhini Infraorder (-formes or -rhini): Platyrrhini Superfamily (-oidea): Ceboidea Family: (-idae): Cebidae Subfamily (-inae): Aotinae Genus: Callicebus Species: moloch [always underline or italicize genus & species names] To see one taxonomy of the living primates, click here IV. The 6 Main Primate Groups A. Lemurs (Madagascar) B. Loris group (Africa, South Asia, & Southeast Asia) C. Tarsiers (Indonesia and the Philippines) D. New World Monkeys (South and Central America) E. Old World Monkeys (Africa, Asia) F. Apes (Africa, Asia) 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 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 Grooming "claw" on pedal digit (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 "park" their infants or cache them in nests while foraging – cathemeral & diurnal species usually carry them Prosimians • Lemuriformes – Lemuridae (lemurs) – Indriidae (indris, avahis, sifakas) – Daubentoniidae (aye-ayes) • Lorisiformes – 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 • 22 or so mostly arboreal species Madagascar Taxonomy: Prosimians (lemur group) Family Cheirogaleidae Mouse & dwarf lemurs Lepilemuridae Sportive lemurs 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) Small Nocturnal Solitary - coprophagy - sluggish Lemuridae "True" lemurs Medium Nocturnal Diurnal Cathemeral Solitary Social monogamy Large multi-male, multifemale groups Indriidae Indri & sifaka & avahi Small (avahi ) Nocturnal (avahi) Solitary (avahi) Medium (sifaka) Dirunal (indri, sifaka) Multi-male, multi-female groups (sifaka) Large (indri ) Social monogamy (indri) - the ring-tailed lemur (Lemur catta) = only significantly terrestrial prosimian - female dominance over males sometimes - 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 multimale, multifemale 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 Tiny Nocturnal Solitary "Special" Features - give birth to twins & cache them in nests - hibernate & accumulate fat seasonally (in tails) Body size Aye Aye (lemur group) Medium Activity Period General Social Pattern "Special" Features 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) Others in the lemur group Body size Activity Period General Social Pattern Small (avahi ) Nocturnal (avahi) Solitary (avahi) Medium (sifaka) Dirunal (indri, sifaka) Multi-male, multifemale groups (sifaka) Large (indri ) Social monogamy (indri) "Special" Features - 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 infraorder : Lemuriformes Anthropoidea Lorisiformes Tarsiformes family: Lemuridae Indriidae Daubentoniidae Loridae Galagonidae Tarsiidae species: lemurs indris avahis sifakas aye-ayes lorises pottos angwantibos galagos tarsiers Tarsiformes 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 Platyrrhini Catarrhini Nostrils Large, separated by wide septum Small, separated by narrow septum Premolars Three (2133 or 2132) Two (2123) Arboreal only Habitat U s e (only capuchin monkey terrestrial at times) Both arboreal & terrestrial forms Diurnal only Diurnal & Nocturnal forms Carry Infants Dorsally Ventrally Infant Care by Adult Ma l e s Sometimes Very Rarely Platyrrhine vs. Catarrhine Dentition: 2133 vs. 2123 (usually) New World Monkeys Ceboidea Family Subfamily Cebidae Common Names Body size General Social Pattern Squirrel monkey Small Large, multi-male, multi-female groups Some Special Features - strictly seasonal breeding _______________________ _______________ Capuchin monkey - large brain _________ - tool use Medium -adaptable & resourceful (like Old World macaques) - weakly prehensile tail Callitrichidae small polyandry tamarins, twins, high paternal investment, reproductive suppression marmosets Atelidae Pithecinae Uakaris & Sakis Medium Socially monogamous Other? Alouattinae Atelinae Howler monkeys Spider monkey, wooly monkey, woolly spider monkey Large Very large One-male, multifemale groups ("harems") Large fusion-fission communities red-faced uakari:sexual selection? Swamp dwellers (hard to study) - LOUD howling - prehensile tail (strong!) - prehensile tails (strong & dextrous!) Interesting social patterns (kind of like chimps and bonobos) Cebids vs. Callitrichids Callitrichids juvenile golden lion tamarin Pygmy marmoset (smallest primate) Callitrichids Golden Lion Tamarins Emperor Tamarin Moustached Tamarin Cotton Top Tamarin Common Marmoset Callitrichids: New Discovery • • • • Wied’s marmoset (Callithrix kuhlii) Usually born as twins Chimerism: an exchange of cells between twins early during embryonic development; result is that most of these monkeys have tissues grown from their twin's cells Germ line too: marmoset fathers can sire their own children and their nephews 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 Cercopithecoidea Cercopithecinae Colobina e "cheek pouch" monkeys leaf monkeys / langurs Buccal pouches Yes No Diet Fruits & other Leaves & other Body Size Medium to Large Large to Very Large Ta i l s Shortish Longish Natal Coat Same as adult or "drab" colors (e.g., black) Often bright, conspicuous colors Continent of Highest Diversity Africa Asia Temperament Active, socially dynamic Couch potatoes? 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 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 fissionfusion communities Large fissionfusion communities Large multimal/ multifemae communitie s 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”