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ANG 6930 Proseminar in Anthropology IIA: Bioanthropology Day 4 ANG 6930 Prof. Connie J. Mulligan Department of Anthropology Mulligan, Copyright 2011 All rights reserved Did everyone get their assignments back? • I’ve graded everything so if you didn’t get something back, that means I didn’t get it • You should have received: – Quiz 1 – Questions/comments from last week – Journal analysis abstract Mulligan, Copyright 2011 All rights reserved Journal analysis • Great abstracts – Average – 19 – Most frequent mistakes • Forgetting to explain your choice of additional journals • Remember to turn your graded abstract in with your final paper Mulligan, Copyright 2011 All rights reserved Quiz #2 Mulligan, Copyright 2011 All rights reserved This week • • Primate evolution, ecology and behavior • • • • Primatology as anthropology Diversity of living primates Primate models for human evolution and behavior Comparison of humans and other primates Reading – – The Human Species, Chpts 5 (Primates), 6 (Primate behavior and ecology), 7 (The human species) Course packet • • • • • • Martin RD. 2002. Primatology as an essential basis for biological anthropology. Evolutionary Anthropology 11:3-6. Strier KB. 2003. Primate behavioral ecology: From ethnography to ethology and back. American Anthropologist 105:16-27. Rieseberg LH and Livingstone K. 2003. Chromosomal speciation in primates. Science 300:267-268. Khaitovich P et al. 2005. Parallel patterns of evolution in the genomes and transcriptomes of humans and chimpanzees. Science 309:1850-1854. Amici et al. 2010. Monkeys and apes: Are their cognitive skills really so different? American Journal of Physical Anthropology 143: 188-197. Judson O. 2008. Wanted: Intelligent aliens, for a research project, New York Times blog Mulligan, Copyright 2011 All rights reserved Next week • Hominoid to hominin • • • • • Dating the ape-human split Australopiths Early hominin subsistence and social organization Origins of genus Homo – Homo erectus, Neanderthals and other archaic humans Reading – The Human Species, Chpts 9 (Primate origins and evolution), 10 (Beginnings of human evolution), 11 (Origin/evolution of genus Homo), 12 (Evolution of archaic humans) – Course packet • • • • • • • • • • “A new kind of ancestor: Ardipithecus unveiled”, Science, 326:36-40. “Candidate human ancestor from South Africa sparks praise and debate”, Science, 328:154155. Klein RG. 2009. Darwin and the recent African origin of modern humans. Proceedings of the National Academy of Sciences 106:16007-16009. “New statistical model moves human evolution back three million years” ScienceDaily, 11/9/2010. Teaford MR and Ungar PS. 2000. Diet and the evolution of the earliest human ancestors. Proceedings of the National Academy of Sciences 97:13506-13511. Conroy GC. 2002. Speciosity in the early Homo lineage: Too many, too few, or just about right? Journal of Human Evolution 43:759-766. Premo LS and Hublin J-J. 2009. Culture, population structure, and low genetic diversity in Pleistocene hominins. Proceedings of the National Academy of Sciences 106:33-37. Hublin JJ. 2009. The origin of Neanderthals. Proceedings of the National Academy of Sciences 106:16022-16027. “Tales of a prehistoric human genome” Science 2009, 323:866-871. Optional (on Sakai) – Noonan JP. Neanderthal genomics and the evolution of modern Mulligan, Copyright 2011 humans. Genome Research 20:547-553. All rights reserved Terminology • Chromosome • Genome • DNA Mulligan, Copyright 2011 All rights reserved Terminology • Chromosome – Molecules in the nucleus that carry genetic information in a linear sequence • Genome – All the genetic information in an organism, i.e. one representative of each chromosome • Nuclear, mitochondrial, chloroplast • DNA – Deoxyribonucleic acid – The molecular basis of heredity • Chromosomes are composed of DNA • The genome is composed of chromosomes Mulligan, Copyright 2011 All rights reserved Terminology • Locus • Gene • Allele Mulligan, Copyright 2011 All rights reserved Terminology • Locus - region of the genome • Gene - DNA sequence that encodes a protein • Allele - one of several alternative forms of a DNA sequence, can be coding or non-coding Mulligan, Copyright 2011 All rights reserved Introduction to Living Primates Mulligan, Copyright 2011 All rights reserved Why Study Primates? • Reasoning by homology – Closely related species tend to be similar in morphology and behavior – Studying nonhuman primates gives insight into human ancestors • Reasoning by analogy – Natural selection leads to similar organisms in similar environments – Studying biological diversity in ecological context sheds light on evolutionary processes Mulligan, Copyright 2011 All rights reserved Why Study Primates? • Primates are our closest relatives – Morphological, biochemical, and behavioral similarity reflects evolutionary relatedness – Reasoning by homology • Primates are a diverse order – Variable ecology, diet, morphology, mating patterns, and social structure – Reasoning by analogy Mulligan, Copyright 2011 All rights reserved Forms of Primate Social Organization Mulligan, Copyright 2011 All rights reserved Solitary • Smallest social group • Females maintain separate home ranges with juvenile offspring • Males may establish own ranges or may defend ranges of several adult females • Adult females and males have infrequent contact, mainly mating • Except orangutans, all solitary primates are prosimians Mulligan, Copyright 2011 All rights reserved Monogamy • One male and one female form long-term pair bond – Show high levels of paternal care of offspring • Share territory with immature offspring • Not very common – Characteristic of gibbons, some small New World monkeys, a few prosimian species Mulligan, Copyright 2011 All rights reserved Polyandry • One female paired with two or more males – May be more than 1 female, but only 1 is reproductively active • Share home range with offspring • Not very common – May occur among some marmosets and tamarins Mulligan, Copyright 2011 All rights reserved Polygyny: One-male • Several adult females, a single resident male, and immature offspring • “Bachelor males” often form all-male groups • Variable dispersal patterns • Common among howlers, langurs, gelada baboons Mulligan, Copyright 2011 All rights reserved Polygyny: Multimale-multifemale • Most common form of primate social organization • Several adult males, adult females, and immature offspring – Most are female-bonded – Males may move between coed groups, remain solitary, or join allmale groups • Lots of variation in terms of size, composition and distribution • Macaques, savanna baboons, some New World monkeys Mulligan, Copyright 2011 All rights reserved Sexual selection infanticide hypothesis • Hrdy (1977) proposed infanticide enhances male reproductive success • Continuing controversy, but growing data – Infanticide reported in ~40 primate species – Researchers have observed >60 infanticidal attacks and many nonlethal attacks – Observed in one-male and multi-male groups • Increasing evidence confirms predictions of hypothesis Mulligan, Copyright 2011 All rights reserved Infanticide linked to changes in male membership and status • C.P. van Schaik reviewed 55 infanticides observed in wild • 47 (85%) followed changes in male residence or dominance rank Mulligan, Copyright 2011 All rights reserved Infanticide shortens interbirth intervals • Lactation generally prevents conception • Cycling resumes quickly if infant dies • Reduces birth intervals by 25-30% in some species • From an evolutionary perspective, new male can more quickly increase the proportion of his offspring in the group Mulligan, Copyright 2011 All rights reserved Males do not kill their own infants • C.P. van Schaik – 40 of 55 infanticides committed by males not in group at conception – 11 of remaining cases by males not seen mating • V. Sommer – genetic relationship impossible or unlikely in 52 of 55 cases • C. Borries et al. – DNA analyses confirm killer father in 22 of 23 cases – Paternity unclear in other case Mulligan, Copyright 2011 All rights reserved Males gain reproductive benefits • C.P. van Schaik – Infanticidal males subsequently mated with mother in 25 of 55 cases – May have mated with mother in 13 other cases • C. Borries et al. – Documented five infanticidal attacks – Four mothers subsequently gave birth – In all cases, DNA identified presumed killer as father Mulligan, Copyright 2011 All rights reserved Unanswered questions • Does infanticide occur more frequently under certain ecological conditions? • Is there a hereditary predisposition to commit infanticide? Mulligan, Copyright 2011 All rights reserved The altruism conundrum • Primates perform altruistic behavior in nature – Grooming, alarm calls, food sharing • Altruistic behaviors decrease individual fitness cannot evolve by natural selection • Altruistic behaviors cannot be favored just because the group benefits Mulligan, Copyright 2011 All rights reserved W.D. Hamilton and Kin Selection • Ordinary selection can favor altruistic behavior if genetic relatives interact selectively – If you increase fitness of a kin, some proportion of your genes have increased chance of survival • Implications – Altruism is limited to kin – Closer kinship encourages more altruism • Closer kin = higher percentage of your genes • rb > c (Hamilton’s rule) r= b= c= average coefficient of relatedness between actor and recipients sum of fitness benefits to individuals affected by behavior fitness cost to individual performing behavior Mulligan, Copyright 2011 All rights reserved Primate Diversity Mulligan, Copyright 2011 All rights reserved Old and new versions of primate phylogeny Old New -Breaks up prosimians -Lemurs more ancestral - ‘Prosimians’ used casually, but not formally Mulligan, Copyright 2011 All rights reserved The primate pattern • No single derived feature shared by all primates • Traits well-suited to arboreal environments – Grasping hands and feet – Binocular (overlapping fields of vision = detail) stereoscopic (image sent from both eyes to both hemispheres = depth) vision • Perceive distance and depth • Controversy over selection of adaptations – Arboreal hypothesis • All traits evolved as arboreal adaptations – Insect predation hypothesis Mulligan, Copyright 2011 All rights reserved The primate pattern, cont • Grasping hands and feet – Opposable big toe, prehensile hands – Flat nails instead of claws, tactile pads – Hindlimb-dominated locomotion • Greater reliance on sight than on smell – Unspecialized olfactory apparatus, reduced in diurnal primates – Eyes moved forward, developed visual sense, color perception Mulligan, Copyright 2011 All rights reserved The primate pattern, cont • Reproduction and life history – Females have small litters, increased birth spacing – Trend toward longer lives and longer periods of gestation, infancy, childhood, and adulthood • Requires more parental care • Brain and behavior – – – – Large brain compared to mammals of similar size More complex brain anatomy Increased learning and behavioral flexibility Cultural transmission of new behaviors • Food washing – began by young macaque (named Imo), taught to all newborn infants although adults did not acquire behavior Mulligan, Copyright 2011 • Not innate behavior, but learned behavior All rights reserved Remember… • None of these traits make primates unique – Example: dolphins have large brains, extended juvenile dependence, and flexible social behavior • Not every primate possesses all these traits – Some prosimians have claws, more developed olfactory senses – Humans have lost opposable big toe Mulligan, Copyright 2011 All rights reserved Mouse Lemur Mulligan, Copyright 2011 All rights reserved Ring-Tailed Lemur Mulligan, Copyright 2011 All rights reserved Ring-Tailed Lemur Mulligan, Copyright 2011 All rights reserved Blue-Eyed Lemur Mulligan, Copyright 2011 All rights reserved Aye-aye Mulligan, Copyright 2011 All rights reserved Sifaka Mulligan, Copyright 2011 All rights reserved Slow Loris Mulligan, Copyright 2011 All rights reserved Tarsier Mulligan, Copyright 2011 All rights reserved Squirrel Monkey Mulligan, Copyright 2011 All rights reserved Spider Monkey Mulligan, Copyright 2011 All rights reserved White-Faced Capuchin Mulligan, Copyright 2011 All rights reserved Great-Nosed Proboscis (Nasalis) Mulligan, Copyright 2011 All rights reserved Zanzibar Red Colobus Mulligan, Copyright 2011 All rights reserved Black-and-White Colobus Mulligan, Copyright 2011 All rights reserved Long-Tailed Macaques Mulligan, Copyright 2011 All rights reserved Bonnet Macaques Mulligan, Copyright 2011 All rights reserved Assam Macaques Mulligan, Copyright 2011 All rights reserved Japanese Macaques Mulligan, Copyright 2011 All rights reserved Japanese Macaques Mulligan, Copyright 2011 All rights reserved Savanna Baboon Mulligan, Copyright 2011 All rights reserved Olive Baboons Mulligan, Copyright 2011 All rights reserved Gelada Baboon Mulligan, Copyright 2011 All rights reserved Mandrill Mulligan, Copyright 2011 All rights reserved White-Handed Gibbon Mulligan, Copyright 2011 All rights reserved Siamang Mulligan, Copyright 2011 All rights reserved Orangutan Mulligan, Copyright 2011 All rights reserved Orangutan Mulligan, Copyright 2011 All rights reserved Gorilla Mulligan, Copyright 2011 All rights reserved Gorilla Mulligan, Copyright 2011 All rights reserved Gorilla Mulligan, Copyright 2011 All rights reserved Bonobos Mulligan, Copyright 2011 All rights reserved Bonobos Mulligan, Copyright 2011 All rights reserved Chimpanzee Mulligan, Copyright 2011 All rights reserved Chimpanzee Mulligan, Copyright 2011 All rights reserved Chimpanzee Mulligan, Copyright 2011 All rights reserved Nonhuman primate biogeography Generally restricted to the tropics Living Fossil only Mulligan, Copyright 2011 All rights reserved Evolutionary morphing • NYCEP – New York Consortium in Evolutionary Primatology • Website to do 3-D reconstructions of extinct primates – Use DNA-based phylogenies and skulls of extant primates to reconstruct skulls of extinct animals • http://graphics.cs.ucdavis.edu/research/EvoMorph Mulligan, Copyright 2011 All rights reserved What’s the difference between monkeys and apes? Mulligan, Copyright 2011 All rights reserved What’s the difference between monkeys and apes? • Monkeys have tails, apes do not • Monkeys have smaller brains relative to body size • Monkey movement is quadrupedal = arms and legs are generally of similar length so their spine is parallel to the ground; apes have longer arms than legs; humans have longer legs than arms • Apes are more closely related to humans than monkeys Mulligan, Copyright 2011 All rights reserved What do these terms mean? • Hominoid • Hominid • Hominin Mulligan, Copyright 2011 All rights reserved What do these terms mean? • Hominoid (Hominoidea) – Apes + humans • Hominid (Hominidae) – Humans and human-like ancestors • Hominin – New term to replace hominid (i.e. humans and human-like ancestors) and hominid = humans + great apes Mulligan, Copyright 2011 All rights reserved Characteristics of Living Humans Distribution and Environment – Humans are the most widely distributed living primate species. Mulligan, Copyright 2011 All rights reserved Characteristics of Living Humans Brain Size and Structure – Absolute brain size is not as useful a measure of intelligence because larger animals tend to have larger brains. – Alternative methods for analyzing brain size and intelligence. Mulligan, Copyright 2011 All rights reserved Characteristics of living humans Figure 11.1: Average brain volume (cubic centimeters) of selected living primates. Mulligan, Copyright 2011 All rights reserved Characteristics of living humans Figure 11.2: Relationship between body weight and brain weight in primates. Mulligan, Copyright 2011 All rights reserved Characteristics of living humans Bipedalism • The physical structure of human beings shows adaptations for upright walking as the normal mode of movement. Mulligan, Copyright 2011 All rights reserved Characteristics of living humans Figure 11.3: The skeletal structure of the feet of a chimpanzee (left) and a modern human (right). Note how the big toe of the human lies parallel to the other toes, facilitating upright walking. Mulligan, Copyright 2011 All rights reserved Characteristics of living humans Figure 11.5: Side view of the skeletons of a chimpanzee (left) and a modern human (right), illustrating the shape and orientation of the spine. (adapted from Campbell, 1985) Mulligan, Copyright 2011 All rights reserved Characteristics of living humans Figure 11.6: The trunk skeletons of a chimpanzee (left) and a modern human (right) drawn to the same size. Note the proportionately shorter and wider pelvis of the human being, reflecting adaptations to upright walking. (adapted from Campbell, 1985). Mulligan, Copyright 2011 All rights reserved Characteristics of living humans Canine Teeth • Human canine teeth are small and do not project beyond the level of the other teeth, serving much the same function as incisor teeth. Mulligan, Copyright 2011 All rights reserved Characteristics of living humans Sex and Reproduction – The fact that humans do not have the estrus cycle has often been cited as a unique aspect of human sexuality. • Exception – orangutans also lack an estrus cycle – Most temperate zone animals only mate during certain seasons and only around the time of ovulation. – Human females cycle throughout the year and often mate at any time during the cycle. • Bonobos mate outside of estrus to some extent Mulligan, Copyright 2011 All rights reserved Characteristics of living humans Sex and Reproduction – Human Childbirth • Single births • Extended period of dependence • Childbirth difficult and complex, requires assistance Mulligan, Copyright 2011 All rights reserved Chimp cognition and behavior • Cooperation and its limitations – http://www.youtube.com/watch?v=3mv8rfJmC Pk&feature=related Mulligan, Copyright 2011 All rights reserved Discussion Mulligan, Copyright 2011 All rights reserved Evolution • What is orthogenesis? Mulligan, Copyright 2011 All rights reserved Social structure • Do you think social structure (monogamy vs multi-male, female dominance, etc) is more genetically determined or ecologically determined? Mulligan, Copyright 2011 All rights reserved Are humans uniquely different from other primates? • Lots of genetic research is looking for something different in humans that signals the unique evolutionary path towards humans – Is there a human gene? – How about a specific signal of natural selection along a human lineage, i.e. positive selection for a gene involved in language or differential expression of a gene? – Differential expression of genes between humans and others? Mulligan, Copyright 2011 All rights reserved Khaitovich et al., 2005, Science • Compared expression levels of genes active in brain, heart, liver, kidney and testis in humans and chimps – How do you interpret these results? Mulligan, Copyright 2011 All rights reserved