Part I

Anthropology
-​ Anthropology is the study of humanity
-​ Anthros means human
-​ Anthropology is made up of
-​ Cultural anthropology
-​ Studies cultural variation among humans
-​ Linguistic anthropology
-​ Studies how humans use language to communicate with one another
-​ Biological anthropology
-​ Studies humans as biological beings
-​ Archaeology
-​ Studies human past based on what’s been left behind
-​ Biological anthropology
-​ Forensic anthropology
-​ Involves applying skeletal analysis and techniques in archaeology to solving
criminal cases
-​ Human biology
-​ studying modern humans
-​ investigating ways that humans adapt to various environments
-​ understanding the process of growth and development
-​ very often anatomists
-​ Paleoanthropology
-​ Studying fossils of humans and their ancestors
-​ Identifying mechanisms of evolution and adaptation
-​ Hominin: Humans, their ancestors up to most recent common ancestor with
chimpanzees
-​ Hominid: Great apes, humans, and their ancestors
-​ More vague than hominins
-​ Bioarchaeology
-​ The study of human remains in archaeological contexts
-​ Paleopathology is the study of ancient diseases
-​ Primatology
-​ Study of non-human primates
-​ Skeletal biology & osteology
-​ Osteology: the study of the skeleton
-​ Anthropometry: the measurement of human proportions
-​ Skeletal biology: development and physiology
-​ Anthropology: sciences or humanities?
-​ Observation→ hypothesis→ data collection and testing→ interpretation→ observation,
etc
-​ Is a cycle
-​ Biocultural perspectives
-​ Culture and biology are intertwined and inseparable
-​ Ex. Getting wisdom teeth out since they aren’t needed anymore
-​ Most modern biological anthropologists conduct research with this in mind
-​ Founders of the Field
-​ Ales Hrdlicka
-​ Anatomist and Founder of Physical Anthropology
-​ Believed that studying the skeletal remains and evolution could help modern
humans
-​ Also incredibly racist
-​ Humans had to have developed in Central Europe
-​ Franz Boas
-​ Founder of the four-field approach of anthropology
-​ Founding academic father to many in anthropology
-​ Strong proponent of humanity as a single species
-​ Emphasized importance of scientific method
-​ Early History of Physical Anthropology
-​ The field began with anatomists seeking to understand human physiology
-​ Craniometry and anthropometry were the main foci of study
-​ Early research led to questionable interpretations
-​ New physical anthropology in mid-20th Century; more holistic and scientific
-​ Semi-transformation in Biological Anthropology rather than the old terminology of
Physical Anthropology
-​ Archaeology
-​ The systematic study of the human past through the materials left behind
-​ History: the study of the human past through written documents
-​ Prehistory: the study of the human past before the development of writing,
using artifacts and other material evidence
-​ Artifacts
-​ Usually not a crystal skull…
-​ Any man-made object
-​ Tools, pottery, bones from animals, bones from humans
-​ Feature- an artifact that cannot be removed
-​ Building, earth mounds, post holes, hearths
-​ Excavation
-​ One level at a time
-​ Pedestal artifacts
-​ Measure and map
-​ Screen all dirt removed
-​ Highly detailed maps
-​ Once excavation has occurred, can never be put back
-​ Must be well-documented
Evolutionary Theory
-​ Culture
-​ “Man’s extrasomatic means of adaptation” - Leslie White 1959
-​ Beyond biology:
-​ Lifeways
-​ Belief
-​ “Material culture”
-​ Altering the landscape
-​ As archaeologists, all we can study are the material culture and landscape and try and
infer the lifeways and beliefs
-​ Evolution is biological
-​ Cultures DO NOT evolve
-​ The correct biological and anthropological definition of evolution: CHANGE in a species OVER
TIME
-​ Not “progress”, “improvement”, or “better than”
Ancestors and How we Know Them
-​ Every culture tells stories of their origins
-​ Epistemes: fundamental cultural ideas of how we know things
-​ In the west, our epistemes are:
-​ Naturalism
-​ Rationalism
-​ Empiricism: everything must be physical, able to measure
-​ Accuracy
-​ These came from the Enlightenment (17th century)
Early Thought
-​ Ancient Greeks credited with first written systematic efforts to understand the natural world
-​ Foundation of current western thought
-​ Aristotle believed things are “fixed” (fixity of species) and never change, and humans are
at the top
-​ Arab scholars preserved the Greek writings
-​ Created more reliable observations and interpretations
Division of the Natural World and Gradual Evolution
-​ Ibn Khaldun (1377 C.E.)
-​ Mineral→ plant→ animal→ monkeys→ human
-​ Gradual change
-​ Al Jahiz (8th Century) and Ibn Miskawayh (10th Century)
-​ Animals come from other animals through gradual changes
-​ Monkeys almost human, thus humans must come from monkeys
-​ Al Jahiz and Nasir Al-Din Al-Tusi (13th Century)
-​ Animals engage in a struggle for existence
-​ Organisms with new features gain advantage and survive
-​ Al Jahiz, Al-Biruni (11th Century) and Ibn Khaldun
 -​ The environment influences new characteristics and thus new species
-​ Causal relationship between environment and appearance
The Roots of Western Science
-​ The Renaissance (14th-16th centuries)
-​ “Rediscovering” the Greeks and Romans (from the Arab scholars)
-​ The study of human anatomy
-​ Belief in a single creation event popular before 19th century
-​ James Ussher’s (17th Century) dating of the world based on biblical references to Adam,
Cain, and Abel- about 5,500 years old
Origin of Species before Darwin
-​ 3 main assumptions before the Enlightenment
-​ No such thing as Extinction
-​ All beings sit in the “Great Chain of Being”
-​ All animals come from Mount Ararat, the site of the landings of Noah’s Ark
-​ By 17th century, all three were in question
-​ Extinction observed first-hand and in fossils
-​ Life as a tree rather than a chain
-​ Biology and geography seemed correlated
Linnaeus and the Natural Scheme of Life
-​ Taxonomy
-​ John Ray
-​ First to use “genus” and “species” to designate types of animals and plants
-​ Carolus Linnaeus
-​ Binomial nomenclature
-​ Systema naturae
-​ Helps make sense of patterns and relationships between organisms
-​ Domain→Kingdom→Phylum→Class→Order→Family→Genus→Species
The Road to the Darwinian Revolution
-​ Georges Cuvier (1769-1832)
-​ Catastrophism
-​ Early forms of life wiped out by event such as Noah’s flood
-​ Believed God purposefully wipes out species
-​ Thomas Rubbert Malthus (1766-1834)
-​ Malthusian Catastrophe
-​ Population growth outpaces resources, population dies off, creating balance
-​ Jean-Baptiste Lamarck (1744-1829)
-​ Theory of the inheritance of acquired characteristics
-​ Concept of “need and use”, didn’t know genes
-​ Relationship between organism and environment
-​ But fundamental change occurs during a single lifetime
-​ ***Lamarckian evolution
The Uniformitarians: Hutton and Lyell
-​ Uniformitarianism: theory that the same gradual geological process we observe today was
operating in the past
-​ James Hutton (1726-1797)
-​ Studied rock formations
-​ Asserted uniformitarianism but did not apply it to the living world
-​ Could not rectify what he observed with his religious beliefs
-​ Charles Lyell (1797-1875)
-​ Key role in convincing public that Earth’s history could only be found in geological
changes
-​ Also a creationist
-​ Friendship with Darwin
-​ Principles of Geology
The Darwinian Revolution
-​ Charles Darwin (1809-1882)
-​ Studied under John Hensolow, a botanist and naturalist
-​ Read travels and natural history of Baron von Humboldt
-​ Voyage on the HMS Beagle
-​ 1831-1836
-​ Charting coasts of South America
-​ The Galapagos
-​ Variations of species
-​ Tortoises
-​ At least thirteen different finch variations
-​ Ornithologist John Gould later sorted them into different species
according to island
-​ Biogeography
-​ Distribution of plants and animals on Earth
-​ Darwin's observations
-​ Oceanic islands hold many species not found elsewhere
-​ Isolated islands lack whole groups of animals, such as large mammals
-​ Distinctive animals and plants resemble close relatives on the mainland even
with a vastly different environment
-​ Darwin’s finches: adaptive radiation of bill types
Refining the Theory of Evolution by Natural Selection
-​ Alfred Russel Wallace
-​ Field biologist in Indonesia
-​ Own version of theory of evolution parallel to Darwin
-​ Darwin published On the Origin of the Species in November of 1859
-​ Darwin’s three observations and two deductions
-​ Observation 1: All organisms have the potential for explosive population growth
-​ Observation 2:Yet populations are roughly stable
-​ Deduction 1: There must be a struggle for existence
 -​ Observation 3: Nature is full of variation
-​ Deduction 2: Some variations are favored while others are not
-​ Refining the Theory of Evolution by Natural Selection
-​ Properties of natural selection
-​ The trait in question must be inherited
-​ The trait in question must show variation between individuals
-​ The environment must exert some pressure on the trait
-​ Fitness- the ability to successfully reproduce
-​ Population- not individuals
-​ Mutation in the genetic sequence- raw source of variation
-​ Post-Darwinian Evolutionary Theory
-​ Darwin did not get rid of Lamarch
-​ Blended inheritance
-​ Mom has blue eyes, dad has brown eyes→ kid would have a mix to blue
and brown eyes
-​ Acquired characteristics to keep variation
-​ Somatic cells and Germ cells are different
-​ “A hen is just an egg’s way of making another egg” -Samuel Butler
-​ Darwin knew no method of inheritance
-​ Mendel and Genetics
-​ Gregor Mendel (1822-1884) observed discrete inheritance
-​ Ex. Kid having blue eyes when parents both present with brown eyes
-​ Work was re-discovered in 1900 and used to explain animal inheritance
-​ New studies in genetics and inheritance led to the Synthetic Theory of Evolution (aka
Evolutionary Synthesis)
-​ Population Genetics
-​ Microevolution - at the genetic level
-​ Macroevolution - at the anatomical level
-​ Modern Evolutionary Synthesis
-​ Combined early 20th Century finding in population genetics with Darwinian evolutionary
thought
-​ Natural selection:
-​ Malthusian Competition
-​ Species Variation
-​ Mutation
-​ Genetic Variation
-​ Mendelian Inheritance
-​ Post-Synthetic Evolution
-​ Clearly genetics and anatomy are not 1-to-1
-​ Background extinctions v. mass extinctions
-​ Diversity in number of species are not equivalent to success- species selection
-​ Debates on HOW and WHEN speciation happens
 -​ Connecting genes to anatomy
-​ Evo-Devo
-​ Seek connection between microevolution and macroevolution
-​ Processes of development lead to phenotypic differentiation
-​ How processes lead to observable patterns
-​ Genetic Toolkit= Homeobox Genes
-​ Econology, Epigenetics, and mechanical stimuli
-​ Biocultural approach
-​ The human species is incredibly “plastic”

The Study of Genetics

-​ Cellular and molecular genetics
-​ Level of basic building blocks (cells) at the most fundamental level of genetic
transmission
-​ Classical and mendelian genetics
-​ Pedigree of related individuals to track how traits are passed from one generation to the
next
-​ Population genetics
-​ Molecular and observable traits for groups of individuals who associate more with one
another than with members of another population
-​ Phylogenetics
-​ Estimating evolutionary relationships between species, usually by constructing treelike
diagrams
The Cell
-​ The basic building block of life
-​ Single-celled organisms
-​ Multicellular organisms
-​ 2 types;
-​ Prokaryotes
-​ Don’t have nucleus→ genetic code floats around
-​ Eukaryotes
-​ Has nucleus consisting of genetic code
-​ Eukaryotic cells
-​ Nuclei
-​ DNA
-​ Gametes
-​ Cytoplasm
-​ Cell membrane
-​ Mitochondria
-​ Ribosomes
-​ Endoplasmic reticulum (ER)
-​ DNA Structure and Function
 -​ Hereditary material must:
-​ Be able to replicate
-​ Be able to make proteins
-​ Coordinate the activity of proteins to produce bodies
-​ Double helix
-​ Nucleotide
-​ “Deoxy” Sugar (C5H10O4)
-​ “Deoxy” since an oxygen molecule has been removed from regular sugar
(C5H10O5)
-​ Phosphate (PO4)
-​ Base
-​ Pyrimidines
-​ Cytosine (Always to G)
-​ Thymine (Always to A)
-​ Purines
-​ Adenine (Always to T)
-​ Guanine (Always to C)
-​ Protein Synthesis and Genes
-​ Proteins
-​ Amino acids
-​ 20 kinds
-​ Polypeptide chains
-​ Compose bone and muscle
-​ Hormones
-​ Enzymes
-​ Cellular function
-​ Genes
-​ Fundamental unit of heredity
-​ The long chain of codons that represent 1 particular protein
-​ Synthesis
-​ Transcription and translation
-​ Codon: 3 nucleotides that code for an amino acid
-​ Introns regulate and make a protein
-​ Exons code for the amino acid
-​ Types of Genes
-​ Structural- produce proteins responsible for morphology of traits (a gene that
codes for a product like enzymes and structural proteins)
-​ Regulatory- control the timing of development and protein synthesis
-​ Homeobox (Hox)- regulate embryological development
From Genotype to Phenotype
-​ Genotype
-​ Set of alleles an organism carries
 -​ Phenotype
-​ Observable feature of an organism that is under genetic influence
-​ Relationship between them is sometimes direct, most often indirect
Alleles
-​ Alleles- Different versions of a gene
-​ Homozygous
-​ Both copies of alleles are same
-​ Heterozygous
-​ Alleles are different
Mendelian Genetics
-​ Gregor mendel (1822-1884)
-​ Blending inheritance (Idea before Mendel)
-​ Experiments in particulate inheritance
-​ Plant breeding
-​ Discovered dichotomous variation
-​ Developed a series of postulates about inheritance
Mendel’s Postulates
-​ Hereditary characteristics are controlled by particulate unit factors that exist in pairs in individual
organisms
-​ When an individual has two different unit factors responsible for a characteristic, only one is
expressed as is said to be dominant to the other, which is said to be recessive
-​ Mendel’s Law of Segregation: During the formation of gametes, the paired unit factors separate,
or segregate, randomly so that each sex cell receives one or the other with equal likelihood
-​ Mendel’s Law of Independent Assortment: During gamete formation, segregating pairs of unit
factors assort independently of each other
Dominance and Recessiveness
-​ Dominance allele
-​ An allele that is always expressed and masks the effect of a recessive allele in a
heterozygote
-​ Recessive allele
-​ An allele whose effect is masked by a dominant allele in a heterozygote
-​ Polygenic trait: a complex genetic trait affected by 2 or more genes and the environment
-​ Pleiotropic trait: a single gene influences many traits
Chromosomes
-​ DNA condenses into chromosomes during cell division
-​ Homologous pairs
-​ 22 pairs of autosome
-​ 1 pair sex chromosomes
-​ 23 pairs of chromosomes total
Rosalind Franklin
-​ Made an essential contribution to the discovery of DNA structure
-​ X-Ray crystallography
Linking and Crossing Over
-​ Linkage and assortment
-​ Linkage: Genes found on the same chromosomes are said to be linked
-​ Crossing Over: genes are exchanged between two chromosomes in pair
-​ Recombination: The result of crossing over
-​ Cross-over important to inheritance occurs during cell division in MEIOSIS
-​ Cross-over in MITOSIS may infrequently occur but usually causes problems
PCR, Mitochondrial DNA, and Ancient DNA
-​ Polymerase chain reaction (PCR)
-​ Taq polymerase extends a strand of DNA
-​ Two primers are placed at each end of the target sequence
-​ Allows for recovery of DNA sequences from miniscule samples
-​ Mitochondrial DNA (mtDNA)
-​ Highly variable and allow for linking to an individual
-​ Has no exchange between maternal and paternal DNA as it is passed through
generations
-​ Passed on only through the mother
-​ Comes from mitochondria floating in the cytoplasm of the egg
-​ Y chromosome acts as another version of mtDNA
-​ Ancient DNA
-​ Bones as old as 100,000 can yield DNA
-​ Possible to recover nuclear DNA sequences
-​ Contamination a concern when PCR technique used
-​ Base pair sequences of Neandertal genome ar about 99.5% identical to our own
Phylogenetics
-​ Not precise reconstructions of evolutionary events, but rather hypotheses
-​ Constructing a phylogenetic tree
-​ Branch
-​ Represents closely-related lineage
-​ Node
-​ Represents separation of any pair of populations
-​ Longer branches and deeper nodes= more changes in DNA
How Evolution Works
-​ Forces of Evolution: Those factors, which occur in natural populations, that cause changes in
gene frequencies over multiple generations
-​ Both adaptive and non adaptive causes
-​ Polymorphism: multiple forms of a trait leading to different phenotypes in a species
Population Genetics
-​ The quantitative study of how genes are distributed both within and across populations
-​ Considers how gene distributions are patterned across time and space
-​ Variation in a population is modified by the forces of evolution
Defining Population
 -​ Population: a group of individuals of the same species who are geographically near enough to
one another that they can breed and produce new generations of individuals (gene pool)
-​ Subspecies: group of local populations that share part of the geographical range of a species and
can be differentiated from other groups based on one or more phenotypic traits
-​ Polytypic species: a species divided into local population that differ by one or more phenotypic
traits
-​ Race: used interchangeably with subspecies in biological sciences; has sociocultural meaning for
humans
-​ Deme: polytypic populations that are defined in terms of genetic composition, such as allele
frequencies
-​ Clade: a group of organisms believed to have evolved from a common ancestor
Microevolution
-​ Changes in the frequency of alleles within a population from one generation to the next
-​ (compare to Macroevolution= the origin of new species)
-​ Modern view of evolution focuses on genetic changes produced by evolutionary forces acting
within populations
Forces of Microevolution
-​ Allele frequencies can change between generations in a population through the following forces
of evolution:
-​ Mutation
-​ Increases diversity
-​ Natural Selection
-​ Decreases diversity
-​ Gene Flow
-​ Increases diversity
-​ Genetic Drift
-​ Decreases diversity
Mutation
-​ Mendel did not know about the biochemical mechanisms of heredity
-​ Knew of spontaneous appearance of variants in a species
-​ Any change in a DNA sequence that becomes established in a daughter cell
-​ Mutations in germ cells can be passed to subsequent generations
-​ Will be present in all cells of the offspring
-​ Could occur during reading
-​ Insertion
-​ Deletion
-​ Could occur from external forces
-​ UV Crosslinking
-​ Sometimes corrected by DNA repair mechanism (ex. enzymes)
-​ Most mutations occur in somatic cells
-​ Only heritable if occurring on gametes
-​ Spontaneous Mutation: a mutation that appears randomly in a family lineage
Point Mutation
-​ DNA sequence incorrectly copied resulting in the change of a single nucleotide base and an error
in protein synthesis
-​ But most point mutations have little impact, since there are 64 DNA triplets that code for 20
amino acids
Missense Mutation
-​ Point mutation that causes a different amino acid in the protein
-​ Could be neutral but could also cause changes in the folding of the protein
Nonsense Mutation
-​ Point mutation creates a codon signaling a stop to the reading
-​ Causes disruption the reading of the gene and creation of the protein
-​ Could be extremely harmful or minor depending on the location of the mutation
Silent/ Synonymous Mutation
-​ Can occur outside of a gene or in the portion of DNA that does not code for a protein (intron)
-​ Silent mutations don’t alter the protein that gets synthesized
Splice Site Mutation
-​ Can neglect to remove an intron or remove a necessary exon
-​ Causes a change in the amino acid sequence
-​ Like Missense, could be neutral but could also cause extreme mis-folding of the protein
Frameshift Mutations
-​ Insertion or deletion mutation (indel) caused by the shifting of the reading frame by not
multiples of three
-​ Extensively change the amino acid sequence
-​ Create new enzyme binding sites or may cause the protein to lose all function
Transposons
-​ Fragments of DNA that can move around the genome
-​ Class 1: Retrotranspons
-​ Transcribed to RNA and then re-inserted (reverse transcribed) in a new location
-​ Class 2: DNA transposons
-​ Directly cut out and re-inserted elsewhere
-​ Major disruptors of genes
-​ Accelerate mutation rate
Judging Mutations
-​ “Bad” mutations (deleterious)
-​ Reduction in the protein’s ability to function
-​ “Neutral” mutations
-​ Common because of redundancy in genetic code
-​ No change in protein form or function or slight changes with no effect on reproduction
-​ “Good” mutations (beneficial)
-​ Increase protein’s ability to function, enhances fitness
-​ Once spread through population, no longer “good”, but “normal” or “wild” type
Natural Selection
 -​ Natural selection is the filter that acts on variation
-​ Phenotypes is physical
-​ Genotype is in the genome
-​ Natural selection operates on the phenotype of an individual organism not the genotype
-​ Changes in gene frequencies
-​ Slow to change over time, populations evolve not individuals
-​ Two kinds of selection: directional and stabilizing
-​ Modes of natural selection
-​ Stabilizing selection
-​ Disruptive selection
-​ Directional selection
-​ Sexual selection
Stabilizing/ Balancing Selection
-​ Maintains a certain phenotype by selecting against deviations from it
-​ Keeps populations uniform
-​ Ex. birth weight of human infants
Disruptive/ Diversifying Selection
-​ Both extremes of a trait are advantageous
-​ Individuals with traits in them idle are selected against
-​ Could lead to high levels of diversity and even speciation
Directional Selection
-​ Drives evolutionary change by selecting for greater or lesser frequency of a given trait in a
population
-​ What you generally think of when you hear “natural selection”
Heterozygous Advantage
-​ Heterozygous genotype has higher fitness than homozygous
-​ Most likely cases where the homozygous recessive is detrimental
-​ Expect that recessive genotype will be lost but persists
-​ Examples:
-​ Sickle Cell Anemia
-​ Cystic Fibrosis
Non-Random Mating
-​ Assortative Mating: selection to mate with those of similar phenotype
-​ Think of dog breeds and the ease/difficult of their mating
-​ Disassortative Mating: selection to mate with those of different genotype to increase diversity
-​ Pheromone selection for complementary immune system
-​ Artificial selection: when humans make the choice for domestic animals
Gene Flow
-​ Movement of genetic material between populations
-​ Migration refers to animals on the move
-​ Admixture: Genetic contributions from other nearby populations
-​ Inbreeding: Reproduction within close populations
 -​ Even small amount of gene flow can mitigate risks
Genetic Drift
-​ Random changes in gene frequency in a population
-​ More impact in smaller populations
-​ Founder effect
-​ Small subset of population cut off from parent gene pool
-​ Ex. The Amish
-​ Ellis-van Creveld (EVC) syndrome and polydactyly more common than general
population
-​ Could be the result of anything
-​ One group with a specific allele reproduces more than another group
Genetic Bottleneck
-​ Form of genetic drift
-​ Rapid reduction in population size, then increase
-​ Only mutations rebuild genetic diversity over a very long time
-​ Founder effect is one type of a bottleneck event
Adaptive Radiation
-​ Once diversity has entered a species, it can quickly fill multiple ecological niches
Taxonomy and Speciation
-​ Homology
-​ Similar features in two related organisms look alike because of a shared evolutionary
history
-​ Analogous traits
-​ Similar traits due to similar use, not due to shared ancestry
-​ Convergent evolution
How Species are Formed
-​ Cladogenesis
-​ When a species branches into two or more new species
-​ Anagenesis
-​ One species evolves into another over time
Types of Speciation Events
-​ Allopatric
-​ Geographic separation triggers speciation between two populations of the same species
-​ Parapatric
-​ Over large geographic area, speciation can occur even with gene flow
-​ Hybrid zones remain and are stable
-​ Sympatric
-​ Ecological factors create more than one phenotype in a single population
-​ No spatial separation necessary
-​ Really only in plants
The Tempo of Speciation
-​ Gradualism
 -​ Slow evolution over vast periods of Earth’s history
-​ Gaps in the Fossil Record?
-​ Fossil record is fragmentary by nature
-​ Macroevolution with major changes over short periods of time
-​ Punctuated equilibrium
-​ Stable with bursts of change

Primates
-​ Mammals are divided into three groups
-​ Prototheria, or monotremes
-​ Lay eggs but nurse their young
-​ Ex. platypuses, echidnas
-​ Metatheria, or marsupials
-​ No placenta
-​ Offspring born in near embryonic state
-​ Ex. kangaroos, koalas, opossums
-​ Eutheria, or placental mammals
-​ Primates
-​ Diversity of Nonhuman Primates
-​ 400+ species of nonhuman primates are recognized
-​ Size and form vary greatly
-​ Body shapes vary tremendously
-​ What Exactly is a Primate?
-​ Mammals with a suite of traits that distinguish them from all others
-​ Grasping hands
-​ Flattened nails
-​ Forward facing eyes
-​ Large brains
-​ High degree of learned behavior
-​ Traditional naming systems
-​ Prosimian-
-​ Lemurs, lorises, galagoes, tarsiers
-​ Anthropoid
-​ Monkeys, apes, hominins
-​ Two suborders of Primate
-​ Strepsirhini
-​ Includes prosimian, except tarsiers
-​ Haplorhini
-​ Tarsiers, monkeys, apes, and humans
-​ Primate Anatomical Traits
-​ Generalized body plan
-​ Many Strepsirhines move by vertical clinging and leaping (VCL)
 -​ Monkeys run and leap along branches rather than swing
-​ Ape’s arm has full range of motion
-​ Petrosal bulla
-​ Skeleton that covers the inner ear
-​ Single bony trait shared by primates that occurs in no other mammalian group
-​ Enclosed bony orbits in the skull
-​ Anatomical adaptation to the importance of vision
-​ May better protect the eye
-​ Prosimians missing posterior wall
-​ Generalized teeth
-​ Infers patterns of behavior and diet
-​ Dental arcade includes four types of teeth
-​ Large Brains
-​ Encephalization
-​ Increase in volume of neocortex
-​ Increased surface area of the brain
-​ Single offspring
-​ Exception is marmosets and tamarins, which give birth to twins
-​ Investment of time and energy in a few babies
-​ Extended ontogeny
-​ Learned behaviors
-​ Many animals live longer than primates
-​ Extended stages in the life cycle
-​ Process of learning to live in a group
-​ Activity patterns
-​ Most haplorhines= diurnal
-​ Many strepsirhines= nocturnal
-​ Sociality
-​ Living in groups
-​ Adaptation by which a primate survives and reproduces
-​ Exception is orangutan
-​ The Nonhuman Primates
-​ Primate families that are anatomically similar are lumped in the same superfamily
-​ Subgroups of families are called subfamilies
-​ The Strepsirhines- Lemurs
-​ Strepsirhines:
-​ Tooth comb
-​ Grooming claws
-​ Lemurs
-​ Found only in Madagascar
-​ Lemuroidea
 -​ Many species now extinct from hunting, including sloth lemurs\Lacked
natural predators
-​ Each of the four species distinct in size and social patterns
-​ The Strepsirhines- Lorises
-​ Lorises
-​ Lorisidae
-​ Galogonidae (bush baby)
-​ Only venomous primate
-​ Communicate vocally and through olfactory methods
-​ Nocturnal
-​ All are slow-moving, deliberate stalkers
-​ Only strepsirhines in Asia and mainland Africa
-​ The Haplorhines
-​ Process full suite of adaptations characterizing living primates
-​ Emphasis on vision
-​ Haplorhines are mostly diurnal
-​ Tarsiers are unique
-​ Most are nocturnal
-​ Traits mixed between prosimian and anthropoid primates
-​ Live in Indonesia
-​ Monogamous pairs
-​ Haplorhines- New World Monkeys
-​ Infraorder Platyrrhini
-​ Refers to the flare shape of the nose
-​ Superfamily Ceboidea
-​ Shared features
-​ Small body size
-​ Three premolar teeth
-​ Arboreality
-​ Prehensile tails
-​ Callitrichids (marmosets and tamarins)
-​ Unique among primates; traits resemble lower mammals
-​ In some species, a polyandrous mating system occurs
-​ Haplorhines- Old World Monkeys
-​ Infraorder Catarrhini
-​ Includes OWM, apes, and hominins
-​ Superfamily Cercopithecoidea
-​ Shared traits
-​ Ischial callosities
-​ Bilophodont molars
-​ Estrus (in some)
-​ Period of sexual receptivity that correspond with ovulation
 -​ Sexual dimorphism
-​ Colobines and Langurs
-​ Leaf-eating monkeys
-​ Diets based on leaves
-​ Able to access food in highly competitive regions
-​ Semi-chambered foregut
-​ Have a distinct paunch
-​ Colobus are in Africa
-​ Langurs are in Asia
Hominoids
-​ Apes
-​ Hylobatidae
-​ Ponginae
-​ Hominane
-​ Shared traits with rest of primates but more extreme
-​ Encephalization
-​ Brachiation
-​ Extended ontogeny
-​ Social complexity in varying degrees
-​ Gibbons
-​ 3 genera; 16 species
-​ Asia, Indonesia
-​ Frugivorous
-​ Most vocal of all nonhuman primates
-​ Mated pairs sing together
-​ Socially monogamous but not necessarily reproductively monogamous
-​ Orangutans
-​ Extremely sexually dimorphic
-​ Bimaturism- Adults take two different forms
-​ 8 year interval between births and reach maturity around 15
-​ Both males and females leave the birth group at maturity
-​ Males are frequently loners holding territory
-​ Females will be included in the territory of a male
-​ Gorillas
-​ Largest primates and extremely sexually dimorphic
-​ Highly cohesive groups
-​ Silverback or blackback males
-​ Varied diets
-​ Mountain gorillas
-​ Lowland gorillas
-​ All prefer to eat fruit but can fall back on fibrous leaves if it is not available
-​ Virunga Volcano groups eat primarily leaves
-​ Chimpanzees
-​ Fission-fusion groups
-​ Males highly social
-​ Form coalitions to hunt and dominate females
-​ Females more independent
-​ To avoid feeding competition
-​ Highly diverse diet
-​ Fruit
-​ Plant products
-​ Termites
-​ Animal meat
-​ Bonobos
-​ Close relatives of chimpanzees
-​ Moderate sexual dimorphism
-​ Live in large, fluid social groupings with strong female bonds
-​ Largely fruit diet, leafy plant material
-​ Have been observed eating meat in some communities
-​ Hypersexual for social interactions
-​ Primate Ecology
-​ Ecology: study of interrelationships of animals, plants, and their physical environment
-​ Jane Goodall, Dian Fossey, Birute Galdikas
-​ Diet
-​ Most primates are herbivores
-​ Frugivores
-​ Folivores
-​ Dietary Constraints
-​ The energy expended to find food== quantity and quality of the food eaten
-​ Primates are color-vision foragers to find the most nutritious fruit
-​ Activity budgets
-​ Time animals spend per activity
-​ Linked to dietary quality
-​ Dietary and Digestive Strategies
-​ Digestive specializations
-​ Foregut of colobine monkeys that allows diet of high-fiber, low-quality leaves
-​ Secondary compounds
-​ Indigestible, toxic chemicals in plant leaves
-​ Defense against leaf eating animals
-​ Diet and Feeding Competition
-​ Feeding competition
-​ Well-defined areas of their habitat to find food and shelter
-​ Intragroup
-​ Intergroup
 -​ Natural food shortages have a severe effect on wild populations
-​ Increase feeding competition
-​ Natural selection may favor individuals that are the best foragers
-​ Territories and Ranges
-​ Defined home ranges
-​ Can be limited or many square kilometers
-​ Must contain all resources needed by a nonhuman social groups
-​ Territory
-​ Defended against other members of the same species
-​ Key resources that nonhuman primate groups are willing to protect
-​ Predation of Primates
-​ Observation of predators is difficult
-​ Stealthy
-​ Nocturnal
-​ Solitary
-​ Small-bodies primates more vulnerable
-​ Behaviors that may have evolved in response to threat of predation
-​ Even humans pose a threat
-​ Primate Communities
-​ Primate communities are integral parts of tropical forest ecosystems
-​ Niche separation
-​ Occurs among all primates that are sympatric
-​ Makes primates highly diverse
-​ Primatology
-​ Studying sociality
-​ Costs and benefits of group living
-​ Models of group living
-​ Proxy for human evolution and social lives
-​ Field Study
-​ Habitat in which species evolved ​
Positional behavior, social interaction
-​ Smaller amount of research can be done
-​ How we Study Primates
-​ Captive study
-​ Close, personal observations
-​ Ability to manipulate the study group
-​ Animals in highly unnatural settings
-​ Semi-free ranging environment
-​ Very large enclosures or small island
-​ Compromise between captivity and natural field study
-​ The Evolution of Primate Social Behavior
-​ All behaviors seen in the wild have proximate causes
 -​ Hunger, fear, sexual urges
-​ Hormonal and/ or physiological reasons to act
-​ Behaviors may be evolved strategies for reproductive success
-​ We’ve already talked about the forces of evolution and how natural selection works
-​ Examine how these forces shape primate phenotype and potentially behavior
-​ The Evolution of Primate Social Behavior
-​ Value of an evolutionary approach is to allow us to test hypotheses regarding fitness
-​ Mating behavior
-​ Physiology
-​ Dominance relationships
-​ Coalition networks
-​ Behavior can be seen as an adaption, phenotype
-​ Social system- the grouping pattern in which a primate species lives
-​ Including its size and composition
-​ Potentially evolved in response to natural and sexual selection pressures
-​ Philopatry
-​ Most female kin living together share a strong incentive to cooperate over food
resources
-​ When females are in fission-fusion situations, male bonds are the most effective way of
controlling females
-​ Social Behavior and Reproductive Asymmetry
-​ Females invest more energy and time in offspring and have lower reproductive potential
-​ Competed over by males
-​ Prioritize obtaining adequate food for themselves and offspring
-​ Female philopatry
-​ Females do not migrate at maturity
-​ Remain and breed within birth group
-​ Male philopatry
-​ Males do not migrate at maturity
-​ Remain and breed within birth
-​ Females in male-philopatric societies may not show a high degree of affiliation
-​ Number of males is dependent on the risk of predation
-​ The degree of parental investment is key in shaping the social system
-​ Males rarely engage in parental caregiving
-​ Exceptions: Tamarins and Marmosets
-​ Male Reproductive Strategies
-​ Dominance
-​ Established early in life through aggressive play
-​ Males not dominant sex in all species
-​ Rarely is there a linear dominance hierarchy
-​ Flexible in various interactions
-​ Cause for dominance debated
 -​ Reproductive success results mixed
-​ More likely for high-ranking individual to be reproductive
-​ Female Reproductive Strategies
-​ Dominance
-​ Small but significant influence
-​ Female non humans typically competed over, but they do not always mate with the male
winner of the competition
-​ Choosiness over mates because of the time invested in offspring
-​ Sexual Receptivity and Signaling
-​ Restricted fertility
-​ Posture
-​ Hanuman Langurs arch their tails over their backs and shake their heads from
side to side
-​ Female features attract mates and may incite males to compete over them
-​ Estrus: During ovulation, a female’s rump may change color, swell, or emit odors to
signal males
-​ Swellings evolved independently at least three times in the primate order
-​ Do no always signal ovulation
-​ May serve to confuse paternity and discourage aggression
-​ Sexual Dimorphism and Society
-​ Access to mates
-​ Competition energy expenditure and injuries
-​ Intensity of competition is reflected in the level of sexual dimorphism in
primates
-​ Multi-female societies are more dimorphic- usually 30-40%
-​ Exception: Orangutans: males are 50% larger than females
-​ Gibbons-single pair-very little sexual dimorphism
-​ The Paradox of Sociality
-​ Group living is an evolved primate adaptation
-​ Benefits:
-​ Improved access to mates
-​ Improved access to food
-​ Protection from predation
-​ Costs
-​ Competition
-​ Enormous energy expense
-​ Sociality means culture
-​ Food
-​ Benefit of exploiting food-finding abilities of others offset by
competition once food is found
-​ Fighting over food, once it’s found
 -​ Females are highly dependent on food availability due to energy
expenditure of reproduction
-​ Predators and Sociality
-​ Avoiding predators
-​ Little direct evidence
-​ Indirect evidence suggests benefits to group living for avoiding predation
-​ Alarm calls
-​ Individual’s chance of becoming the victim lowered
-​ Types of Primate Societies
-​ Social system
-​ The grouping pattern in which a primate species lives, including its size and
composition evolved in response to natural and sexual selection pressures
-​ Solitary
-​ Monogamy
-​ Polygamy
-​ Polygyny
-​ Polyandry
-​ Polygynandry
Solitary
-​ Strepsirhines and Orangutans
-​ Females occupy individual territories with dependent offspring
-​ Males occupy territory overlapping several female territories and decide whether to
allow access to transient males
-​ Males use scent markers to define territories (strepsirhines)
-​ Most solitary primates are also monogamous
Monogamy
-​ Male and female live in a pair bond for an extended period of time
-​ Gibbons
-​ No strict reproductive monogamy
-​ Sometimes they mate outside of the pair bond
-​ Best understood as a female reproductive strategy
-​ Fewer aspects of male competition
Polygny is Multi-female
-​ One-male polygyny
-​ Formerly harem, but that term implies control over females
-​ One male lives with many females he hopes to monopolize
-​ Gelada baboons- females are bonded to each other and the male hangs around hoping
to have access to them
-​ Gorillas-females may migrate between silverbacks. Males have anxiety that the females
will leave for another silverback
-​ Extra males typically reside in all-male “bachelor” groups
-​ Hanuman langurs
 -​ Commonly compete for takeovers and practice infantice (kill offspring of
previous males)
Multimale polygyny
-​ Reduce extraneous competition but maintain largest share of females through dominance
-​ Dominance hierarchy to compete for priority of access to females
-​ Seasonal breeding leads to intense competition
-​ Some species exhibit both one-and multi-male polygyny
-​ Sexual dimorphism
Fission-fusion polygyny/ polygynandry
-​ Chimps, bonobos, spider monkeys
-​ Temporary associations of individuals- communities
-​ Based on foraging parties
-​ Only stable unit is a female and her offspring (nuclear family)
-​ May be an evolved response to reliance on ripe fruit
-​ Seasonal distribution and daily variation in fruit
-​ Females forage for increased access to fruit
-​ Males form bonds with one another to control access to females
Polyandry is multi-male
-​ Polyandry
-​ Marmosets and tamarins
-​ One female with multiple males
-​ Poorly understood and rare in primates
-​ Males bond together and help females rear offspring
-​ May be reproductively monogamous but socially polyandrous
Fossil Context & Geologic Time
-​ Setting the Stage
-​ For understanding human evolution
-​ Looking at geology
-​ Paleontology
-​ Understanding fossilization
-​ Explore conditions on earth during the Cenozoic
-​ Look at some of the earliest primates
-​ Paleontology
-​ The study of extinct organisms based on their fossilized remains
-​ Taphonomy
-​ The study of what happens to the remains of an animal from the time of its
death to the time of discovery
-​ Process of becoming a fossil
-​ Burial
-​ Carcass covered with sediment
-​ Interrupts decomposition
-​ Basics: How to Become a Fossil
 -​ Petrification
-​ Skeletal remains absorb surrounding minerals that eventually replace the
organism’s inorganic tissues
-​ Most fossilized remains are of skeletal parts but occasionally skin; hair, or plant parts are
preserved
-​ Trace fossils
-​ Such as tracks
-​ Coprolites
-​ Fossilized feces
-​ The Importance of Context
-​ Without context, a fossil cannot be assessed for age or environment
-​ To a paleoanthropologist- context is everything​
Established by
-​ Stratigraphy
-​ Geologic Time Scale
-​ Provenience- the origin or original source (as of a fossil)
-​ Principles of Geology
-​ The principle of original horizontality
-​ Layers are originally parallel to Earth’s gravitational field
-​ The principle of superposition
-​ Older layers are laid down first and covered by younger layers
-​ The principle of cross-cutting relationships
-​ A geological feature must exist before another feature can cut across or through
it
-​ Stratigraphy
-​ Strata are layers in rock
-​ Stratigraphy Tapho
-​ The study of the order of rock layers and the sequence of events they reflect
-​ The principle of faunal success
-​ Successive layers contain certain types of faunal communities with predictable
patterns
-​ Index fossils typify the animalism in a layer
-​ The Geologic Time Scale (GTS)
-​ GTS is categories of time into which the earth’s history is usually divided
-​ The earth is about 4.5 billion years old
-​ Human and primate evolution spans the last 65 million years (Cenozoic Era)
-​ The GTS is divided into eons, eras, periods, and epochs
-​ Vastly more time is represented by the Hadean, Archean, and Proterozoic (about
4 billion years) than by all of the later periods, which span only the last 540
million years
-​ Boundaries are points in the time scale where large shifts are evident in the
geologic column
-​ The Anthropocene
-​ The Holocene
-​ Began approx. 10kya
-​ Beginning of warming after last major ice age
-​ Human driven warming
-​ Faster than natural processes
-​ Significantly increased in the last 100 years
-​ Anthropocene
-​ Human driven mass extinctions and warming temperatures
-​ Three possible boundaries: Agriculture (5-8kya), Industrial Revolution (1760),
Nuclear Age (1950)
-​ Types of Fossils
-​ Plants
-​ Petrified wood
-​ Animal remains
-​ Amber
-​ Asphalt
-​ Form of crude oil often called tar
-​ Igneous rock
-​ Trace fossils
-​ How do we know the age?
-​ Relative dating techniques
-​ Age of a fossil only in comparison to other materials found above and below it
-​ Calibrated Relative Dating Techniques
-​ Regular processes calibrated to a chronological scale if conditions are known
-​ Chronometric Dating Techniques
-​ Estimate antiquity of an object in years before present (BP) with precision
-​ Relative Dating Techniques
-​ Lithostratigraphy
-​ The study of geologic deposits and their formation, relationships, and relative
time relationships
-​ Based on their lithologic (rock) properties
-​ Characteristics of rocks correlate across regions
-​ Teprhostratigraphy
-​ Identification of volcanic ash by its chemical fingerprint
-​ Variance of lithostratigraphy
-​ Biostratigraphy
-​ Principle of faunal succession
-​ Index fauna
-​ Rodents often good indicators
-​ Chemical techniques within sites
-​ Analysis of fluorine, uranium, and nitrogen content of the fossils themselves
 -​Calibrated Relative Dating
-​ Geomagnetic polarity
-​ Time scale composed of the sequence of paleomagnetic orientations of strata
through time
-​ Reversed polarity
-​ Dendrochronology
-​ Tree rings correspond to rate of growth over a particular period
-​ Those can be matched up by year to get calibrations on the date the tree was
cut down
-​ Up to 14kya in Northern Hemisphere
-​ Chronometric Data Techniques
-​ Radiometric dating
-​ Relies on natural, clocklike decay of an element or its variant form (isotopes)
-​ Half-life is the amount of time it takes for one-half of the original amount to decay
-​ Parent is the original radioactive isotope. Daughter is the product
-​ Isotopes
-​ Radiocarbon dating
-​ Primary technique for estimating antiquity from Pleistocene through the present
-​ All living organisms contain carbon
-​ Decay rate of 14C is half life of 5,730 years
-​ Limited to objects younger than 40,000 years
-​ Electron Trap Techniques
-​ Measure the effect of exposure to radioactivity
-​ Thermoluminescence (TL)
-​ Optically stimulated luminescence (OSL)
-​ Electron spin resonance (ESR)
-​ Amino Acid Racemization (AAR)
-​ Amino acids have two forms (isomers)
-​ In life, all amino acids are L-form
-​ After death, they flip to D-form at predictable rates
-​ Can be used to more than a million years ago (Mya)
-​ Dependent on temperature
-​ Must know relative temperature fluctuations
Mammals and Dinosaurs
-​ Placental mammals originated in Mesozoic (~250 Ma0
-​ Small, nocturnal, slow, and quiet
-​ Extinction of dinosaurs ~65 Ma
-​ Opened ecological niches to be filled by mammals
-​ Explosion of diversity in Paleocene, suddenly reduced in Eocene
Earth in the Cenozoic
-​ Pangea begins to break up around 200 million years ago
-​ The earth was divided into two major land masses during Jurassic
 -​ Laurasia
-​ Present-day North America, Europe, and Asia
-​ Gondwanaland
-​ Africa, South America, Antarctica, and Australia/ India
-​ We can reconstruct environmental conditions from several kinds of geological and biological
evidence
Environment in the Cenozoic
-​ Vegetation
-​ Compare ancient environments to today
-​ Plant macrofossils
-​ Fossil pollens
-​ Presence of Phytoliths, or opaline silica bodies
-​ Isotopes
-​ Global temperatures​
Plant types
-​ Migrations
-​ Animal bones can als be used to infer local environments
-​ Animal communities
-​ Preferred habitats of past animals
-​ Community of animals at a site, not single species
-​ Dietary and locomotor adaptations
-​ Stable carbon isotope ratios in teeth and soil
-​ Reconstruct vegetation in a region
-​ Vegetation map built from isotopes in teeth of fossil animals
-​ Organic carbon in soils comes from local plants
-​ Gives a picture of whether area was open or wooded
Climate Change and Early Primates
-​ Drastic environmental changes around the Oligocene
-​ Absence of large prey animals favored small mammals
-​ Enter small, primate-like mammals
Why Primates (euprimates)?
-​ Difficult to create an evolutionary scenario
-​ Suite of features rather than single feature
-​ 3 Hypotheses
-​ Arboreal
-​ Visual Predation
-​ Angiosperm-Primate Coevolution
Arboreal Hypothesis
-​ Many features evolved to improve locomotion in the trees
-​ Grasping hands and feet
-​ Flexible joints
-​ Reduced olfaction and increased vision
 -​ Binocular vision
-​ Evidence:
-​ Extend primates are arboreal
-​ More “primitive” primates are arboreal
Visual Predation Hypothesis
-​ Developed in the 60s and 70s to counter Arboreal hypothesis
-​ Other arboreal animals don’t have primate traits (e.g., squirrels)
-​ Compared non-arboreal animals to primates
-​ Animals that share trait include predators
-​ Vision to judge distance
-​ Grasping to catch insects
Angiosperm-Primate Coevolution
-​ But many primates today are not predatory
-​ Primates adapted to be on small branches, but insects not there
-​ Counter- Evidence to Predation:
-​ Most insects caught on ground
-​ Predatory primate suse more hearing
-​ Most fossils seem omnivorous
-​ Fruit-seeking behavior
-​ Fruit of angiosperms (flowering) on branches
-​ Grasping for better hold on marginal branches
-​ Depth perception and color vision
-​ Primates evolved during angiosperm revolution (approx 65 Ma)
-​ Better fruit foragers= better seed dispersal
-​ Symbiotic coevolution
Adding Complexity
-​ Parallel traits between Opossums and Primates
-​ Opossums forage fruit AND prey on insects
-​ Insects also attracted to fruits and flowers
-​ Convenient double meal
-​ Other hypotheses have been proposed
-​ Maybe no single scenario accounts for primates
Plesiadapiforms
-​ Mammalian order or suborder appearing in the Paleocene Epoch
-​ Primate-like
-​ Similar molars
-​ Grasping hands
-​ But also:
-​ Unusual anterior teeth
-​ Claws, not nails
-​ No evidence of postorbital bar
-​ No evidence of auditory bulla
-​ Why are they considered ancestral?
-​ Originally, treeshrews thought to be primates
-​ Resemble Plesiadapiforms
-​ Closer to dermopterans
-​ Closest living relative to primate order
-​ Hands and feet more like primates than dermopterans
-​ Even have petrosal bulla
-​ Maybe some genera closer to primates
True Primates of the Eocene
-​ Two main superfamilies
-​ Beginning of the Eocene, declined during the Oligocene
-​ Adapoidea
-​ Omomyidae
-​ Adapoids (strepsirhine ancestors)
-​ Small to medium sized
-​ Slow moving arboreal quadrupeds
-​ Diurnal
-​ Diet of fruit and leaves
-​ Omomyoids (haplorhine ancestors)
-​ More denver than adapoids
-​ Diets of insects and fruit
-​ Nocturnal
-​ Arboreal quadrupeds
-​ The Strepsirhine-Haplorhine Split
-​ Adapoids and Omomyouids split over available food resources
-​ Last common ancestor of all primates was 63 Mya
-​ Omomyoids focused on fruit and insects, developing short snouts
-​ Adapoids ate fruits and leaves, relying on smell, developing long snouts
-​ What Favored the Origin of Anthropoids?
-​ Changes to mandible and eye orbits allowed for chewing toucher foods and better
protection of the eyes
-​ Increase in body size cause they could eat more
-​ Stereoscopic vision
-​ More leave based folivorous diets
-​ Evolution of Tarsiers
-​ Current hypothesis suggest tarsiers developed from Omomyoids
-​ Earliest split probably occurred 50 Ma
-​ Ancestors done evolving by the Eocene
-​ Hypotheses for Anthropoid Evolution
-​ As always, 3 differing hypotheses:
-​ Adapoing Origin
-​ Has been most rejected
 -​ Omomyoid Origin
-​ Develop postorbital closure independently in anthropoids and tarsiers
-​ Tarsier Origin
-​ Genetic data suggests tarsiers and anthropoids are sister groups
-​ Omomyoids too ancient for DNA
-​ New World Monkeys
-​ Molecular evidence for NWM and OWM divergence about 40 MYA
-​ Apidium-like ancestor may have “rafted” over to South America from Africa
-​ Three premolars rather than two
-​ Middle Miocene shows a rich fossil record of NWM
-​ The Divergence of Monkeys and Apes
-​ Evolution of brachiation
-​ Early middle Miocene forests were widespread
-​ Knuckle-walking developed in African apes
-​ Dental evidence
-​ A dietary shift with apes eating more fruits
-​ Apes across the East
-​ Apes appear to have developed in Africa
-​ Middle-Late Miocene had lang bridge between Africa and Eurasia (16 Ma)
-​ Earliest apes were called dental apes
-​ Typical Y-5 molar pattern
-​ Lacking a tail
-​ Otherwise very monkeylike
-​ Late Miocene Apes
-​ Gigantopithecus
-​ Largest primate that ever lived
-​ Ouranopithecus
-​ Massive brow ridge, resembling a gorilla
-​ Dryopithecus
-​ Primitive anatomy like Proconsul and gibbons
-​ Likely closer to apes