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Speciation
A species diverging into 2 distinct lineages, acts as a barrier to reproduction
What does lack of gene flow cause
allele frequencies in isolated populations can diverge = speciation
Why do populations diverge
Natural selection, genetic drift, mutations
Phylogeny
The evolutionary history of a lineage/lineages, similar to a family tree
Species
An evolutionarily independent population or group of populations
Biological species concept
Reproductive isolation is the method of identifying species
Mechanisms that stop gene flow
Pre and Postzygotic isolation
Prezygotic isolation
Individuals of different species are prevented from mating successfully
Postzygotic isolation
Hybrid offspring do not survive/are sterile
Pollinator isolation
Floral traits are adapted to specific pollinators, ensuring only conspecific pollen (from the same species) is transferred
Hybridization
Some closely related species are able to interbreed and reproduce
Is hybridization favoured?
No, natural selection acts against hybrids
What does hybridization signify
Reproductive isolation has not completely evolved between the 2 species since they are able to reproduce
Advantages of Biological species concept
Reproductive isolation signifies evolutionary independence
Disadvantages of Biological Species Concept
Cannot be evaluated in fossils species or species that reproduce asexually and difficult to assess if populations do not overlap geographically
Morphological species concept
Species are those that look alike
Advantages of Morphological Species Concept
Useful for lack of data on extent of gene flow, applicable to sexual, asexual and fossil species
Disadvantages of Morphological Species Concept
A single polymorphic species may be incorrectly classified as more than 1 species, states that members of the same species all look alike, can be disproved (Ex. butterflies), and traits used to distinguish species are subjective
Ecological Species Concept
Species are organisms that exploit the same resources, have similar environmental tolerances and face the same predators/parasites
Advantages of Ecological Species Concept
Useful for identifying asexually reproducing species
Disadvantages of Ecological Species Concept
Cannot be defined by reproductive isolation from other species - explained by adaptations to ecological conditions
Phylogenetic species concept
Smallest monophyletic group on a phylogenetic tree indicates a species, based on synapomorphies
Advantages of Phylogenetic species concept
Applicable to any type of population (Ex. fossil, asexual, sexual) and logical
Disadvantages of Phylogenetic Species concept
Phylogenies are only available for a small amount of populations, Independent Lineage Sorting, and leads to recognition of many more species compared to other species concepts
What assumptions does the Phylogenetic Species Concept make
All species are related by common ancestry
Independent Lineage Sorting
When ancestral genetic variation is randomly distributed into descendant species, causing gene trees to differ from the actual species tree (Ex. A -> B, C can turn into B -> A, C: makes AC look more closely related than BC)
Gene tree discordance
Gene trees do not always match species trees
Subspecies
Populations that live in discrete geographical areas, have distinguishing features, can interbreed if geographical barriers are removed
Synapomorphies
Homologous traits found in common ancestors and descendants but missing in distantly related ancestors (Ex. Genetic synapomorphy)
Dusky Seaside Sparrow
Had 6 subspecies that formed 2 monophyletic groups
1. Atlantic Coast: Genetically indistinguishable from Seaside Sparrow’s
2. Gulf Coast: Genetically distinguishable from Seaside Sparrow’s
Eventually the Seaside Sparrow went extinct as revitalization attempts failed
Allopatry
Populations that live in different areas
Allopatric Speciation
Speciation through geographic isolation
Types of geographic isolation
Dispersal & Vicariance
Dispersal
Individuals colonize a new area (Ex. Genetic drift)
Vicariance
A geographic barrier arises and splits a population into 2 (Ex. Uplifting mountain range)
Symaptry
Populations that live in the same geographic area
Sympatric speciation
Speciation among species inhabiting same geographic area
What can cause sympatric speciation
External & Internal events, Polyploidy
External events
Sympatry: Disruptive selection based on ecological niches or mate preferences
Internal events
Chromosomal mutations
Polyploidy
An error in meiosis or mitosis results in more than 2 sets of chromosomes, common in plants
Why is polyploidy common in plants
Many diploid plants are closely related to polyploids, polyploids have higher levels of heterozygosity and can tolerate higher levels of self-fertilization (not as affected by inbreeding depression as diploids), and genes on duplicated chromosomes can diverge independently = genetic variation
Autopolyploid
Offspring with a mutation that doubles the chromosome number = all chromosomes are from the same species
Autopolyploid Tetraploids
Arise from self-fertilization of gametes that were produced due to chromosomal error
Allopolyploid
Offspring when parents are of different species and mating results in non-sterile offspring = 2 copies of chromosomes from each parent
Allopolyploid Tetraploids
Arise due to self-fertilization of gametes that were produced as a result of hybridization between 2 species
Tetraploid reproduction
can reproduce with other tetraploids but not diploids
How does polyploidy lead to speciation
Through immediate post-zygotic reproductive isolation from their diploid ancestors
Instantaneous speciation
When 2 species hybridize and the offspring are reproductively isolated from both parental species, may be able to reproduce with other hybrid species
Biogeography
How species and populations are geographically distributed
Homology
Traits shared due to similar ancestry
Homoplasy
Traits that are similar for reasons other than common ancestry (Ex. Convergent evolution)
Evolution of eyes
Originally thought to be convergent due to vast differences in function and morphology between animals
Hypotheses of Eumetazoa eyes
1. Originally eyeless and descendents convergently developed eyes (convergence)
2. Had simple photoreceptor eye spots which were developed into complex eyes (parallelism)
3. Had the primitive makings of an eye and ancestors derived from this (homology)
Adaptive radiation
Rapid production of many descendant species from a single lineage
Criteria for adaptive radiation
Part of a monophyletic group, speciated rapidly, diversified ecologically into many niches
How does adaptive radiation occur
Extrinsic & Intrinsic factors, Ecological opportunity
Extrinsic factors
New favourable conditions or access to new environment (colonization)
Intrinsic factors
Adaptive radiation: Evolution of morphological, physiological or behavioural traits
How does ecological opportunity drive adaptive radiation
The availability of more or new types of resource makes it more likely for species to diverge
Escape-and-radiate coevolution hypothesis
A plant lineage evolves a new defense to escape herbivores and radiates into new species and a herbivore evolves to overcome the defense, radiating into new niches = coevolution
Cambrian explosion
All organisms were unicellular until animals became larger and more complex
Hypotheses of what triggered the Cambrian explosion
1. Higher Oxygen levels: efficient aerobic respiration
2. Evolution of predation; drove morphological divergence among prey (selection for shells, rapid movement, other defenses)
3. New niches beget more: Mass migration off of ocean floor = exploit land resources = new niches
4. New genes, new bodies: Development of Hox genes in animals made it possible for larger more complex bodies to evolve
Early animal fossils (before cambrian explosion)
Microfossils, less than 1mm, first macroscopic fossils included sponges, jellyfish, fossilized tracks, most likely burrowers, floating or immobile feeders
Cambrian fossils
Included wormlike animals, anthropods, mollusks, swam, walked, ran, more mobility overall, filled many ecological niches (ex. Predator, scavenger)
The Burgess Shale Reef
Slit sifted into buried animals preventing animal scavenging and bacterial decomposition = well preserved fossils (Ex. antennae, eyes, digestive tracts)
How did the Burgess Shale Reef provide evidence of predation
Contained fossils with gut contents containing fragments of shelled prey + fossils with healed damage resembling modern predators
Hominids
Great apes, large bodied, long arms, short legs, no tail, monophyletic group that comprise Homo Sapiens + 20 extinct bipedal relatives (Ex. neanderthals)
WHO DA ONLY BIPEDAL APES
Humans! Bipedalism is the synapomorphy that defines hominins
4 general groups of hominid species
Oldest known hominin: Ardipithecus ramidus
Comibation of tree climbing and bidepal activity
1. Gracile australopithecines
2. Robust australopithecines
1&2 small bipeds with small teeth
Evolved big teeth to eat plants
3. Early Homo
Larger braincase and reduced cheek teeth
4. Recent Homo
Even larger braincase
Started to see the use of fire to cook food, used stone utensils
Neanderthals
Last hominin species to live with our species, heavily built, large brains, low foreheads, powerful jaws, large browridges, small chins
First ancient-human hybrid
small piece of bone found in Siberia indicating 40% Neanderthal DNA and 40% Denisovan
Homo sapiens
Reduced browridges, flat faces, largest braincases, larger brain advantage = efficient hunting, tool use, communication, social lives (Trade-off: Energy expensive)
Hominin family tree
no linear progression from one species to another, branches off each other
WHO HAS DA LARGEST BRAINS?
Species in Homo genus have extremely large brains relative to overall body size!
What does the evolution of larger brains indicate
Support a natural selection hypothesis for the ability to reason and communicate, triggered by increased tool and language use = larger brains
Out-of-Africa hypothesis (relative to H. sapiens)
H. sapiens evolved its distinctive traits in Africa and dispersed throughout the world
Mechanism of Out-of-Africa hypothesis
First lineage to branch off led to descendant populations that live in Africa today (highest genetic diversity), subsequent branching = residents of central Asia, Europe, etc. 2 waves of migration: Wave 1 = Australia, Wave 2 = Europe and Mainland Asia
Evidence of H. sapiens interbreeding
1 - 4% of genome of indigenous Europeans and Asians are derived from Neanderthals & 5% of genome of Aboriginal Australians is Denisovian
Why did we evolve bipedalism?
1. Lower surface area exposed to sun, risk of heat stress reduced
2. Hands free to carry food or tools to forage more efficiently
3. Long-distance migration, follow migrating herds
Major modifications to H. sapiens
Position of foramen magnum (spine connection): enters skull from below
Shape of pelvis: Bowl-shaped to support upper body and provide balance
Feet: Lost opposable great toe, larger heel, arched food to propel forward
Large brain: Modern brain development continues until age 20
Evolution of language
No fossil evidence, potential link between toolmaking and language development, related to FOXP2 (language gene)
FOXP2
Essential for human speech, linked to vocalization in mice, song for bids, other verbal comm, Hyoid bone position allowed tongue movements for speech
Evolution of human altruism and cooperation
Evolved through fitness feedbacks in small groups, punishment of cheaters, cooperative hunting
How does the fossil record support the evolution of human cooperation
Visible injuries due to hunting, healing from wounds that required care from others
Do we evolve today?
Yas, coevolution with pathogens, spread of lactose tolerance mutation
Genetic variation in humans
Greater genetic variation within groups than between groups, any 2 humans differ by ~0.5% of genome on average
Examples of human evolutionary traits
Skin pigmentation, Skin cancer, Resistance to malaria, & Lactase Persistence
Skin pigmentation evolution
Over 350 associated genes, natural selection favours more melanin in regions with high UV radiation (Ex. Equator), Light pigmentation = Vitamin D synthesis in regions with lower UV Darker pigment = Protective in regions with high UV
Fitness trade-off of pigmentation
UV breaks down folate and damages DNA but is essential for vitamin D production
Skin cancer types
Melanoma and Non-melanoma
Melanoma
High mortality rate, rare
Non-melanoma
Common, slow progression, rarely metastasizes, low mortality rate
Folate
An essential nutrient that must be obtained by diet, necessary for DNA repair and synthesis, deficiencies decrease chances of successful reproduction. UV radiation causes breakdown in blood folate
Vitamin D
Regulates immune function, protects from pathogens, can be both obtained and synthesized
Resistance to malaria
Deadly infection, targets young children, pregnant women, selection favours modifications of RBC’s that reduce success of parasite (Heterozygote advantage = protection against malaria)
Lactase persistance (pre-evolution)
Milk is a primary food source for young mammals. Nutrient rich: Lipids, proteins, minerals, etc…Digested via lactase during weaning, lactase production is halted post-weaning, lose ability to digest lactose by age 2
Lactase persistance (post-evolution)
Pastoralism: Domestication of cattle, goats, sheep, other milk-producing farm animals. Selection on these populations caused lactase production to be maintained into adulthood
Population
Group of individuals of the same species living in the same area at the same time
Features of a population
Size, density and range