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biogeography (Alfred Russell Wallace)
Similar species tend to be found geographically close together.
Associated with Darwin as well.
common ancestry (Darwin)
The idea that if you trace back the lineages of living species far enough back in time, those species will converge to shared, common ancestors.
homologies (Darwin)
Structures that have deep, underlying similarities between species.
Homologous traits
Same structure, different function. Goes along with homologies and common ancestry studied by Darwin.
nested structure
If taxonomic groups shared common ancestry at different times in the past, it is expected that taxonomies would be structured with groups within groups: multiple genera within each family, multiple families within each order, etc. This nesting pattern is consistent with the idea of a diversifying process, beginning from an ancestor and radiating like the branches of a tree that don’t overlap.
What evidence did Darwin use to justify his theory of common ancestry?
Homologies, transitional fossils, and nested structures of taxonomies
polymorphic
The existence of two or more distinct physical forms, phenotypes, or genetic variants within the same population of a species
Ex. White flower population producing a purple flower (genetic variation). This can happen via gene mutation.
When more than one allele is found at a given locus in a population
evolution
Change in the genetic composition of the population, and it happens when you have a polymorphic population.
fixed or fixation
The change in frequency of a polymorphic population, when variants in a population become extinct.
The loss of all variants except one from a population
An allele that has a frequency of 1.0 in a population
natural selection (Darwin)
Implies that whichever genetic variants make an organism better equipped for the pressures of its environment will tend to increase in frequency in populations over time. Therefore, populations should get better adapted to their environments over time and fixation will occur for these traits. Also, the process of allele frequency change due to fitness variation among genotypes at that locus
transition fossil
Fossil taxa that have some, but not all, of the derived traits of a living group
What is evolution?
Descent with modification that has a pattern (recorded in anatomy, fossils, embryos, genomes, etc.) and process of change (that is observable and testable). It also constitutes a change in the genetic composition of a population over time.
Why do we study evolution?
Unifies biology
It brings together fields like genetics, developmental biology, molecular and cell biology, paleontology, etc.
Practical applications
Combating disease and predicting the spread of pathogens
Fighting drug resistance in bacterial strains
Improving agriculture through development of pest-resistant crops
Supports conservation efforts
Aristotle
Species were fixed (i.e. an elephant is an elephant forever)
He arranged them on scala naturae which created a hierarchy of life forms (“higher” and “lower” species”
He believed that geography changes (water forms change over time and so does land)
William Paley
Natural Theology is what he is known for
He had this idea that a watch implies a watchmaker in the same way that something is complex because of its creator
Life is static but it was created by an entity
Naturally occurring complex structures (like an eye) are comparable to human inventions (like a telescope)
Carolus Linnaeus
Created the binomial classification system (…Order → Family → Genus → Species) in Systema Naturae
He separated organisms into three kingdoms: mineral (geology), vegetable (botany), and animal (zoology)
“Nature does not proceed by leaps and bounds” states that there are bigger differences for a species outside of its genus
He saw species as part of the divine plan, but that Earth and species were unchanging
What was Carolus Linnaeus’s classification system in order?
Imperium (Empire) - everything
Regnum (Kingdom) - animal, plant, mineral
Classis (Class) - in the animal kingdom, there were six: mammals, birds, amphibians, fishes, insects and worms
Ordo (Order) - subdivisions of class
Genus - subdivision of order
Species - subdivision of genus
Varietas (Variety) - species varieties or “sub-species”
James Hutton
Father of modern geology
Proposed uniformitarianism
He believed that natural laws that apply today must also have applied in the past (as in life is not really changing), geological “deep time” (which is geological timescale of Earth’s history that proves the Earth is really really old), and that catastrophic/supernatural events are not required to explain the history of the Earth.
Uniformitarianism (James Hutton)
Changes in the Earth’s crust have resulted from the action of continuous and gradually acting processes, such as erosion and sedimentation
Thomas Malthus
Economist that tried answering the question why people go to war
He created a curve (featured) that shows populations grow much faster that resources, and once that at a certain threshold (after the two lines cross), available resources will limit population growth
This curve later ties in with Darwin’s idea of natural selection, in that species produce more offspring than their environment can support (differential survival and reproduction)

Jean Leopold Nicolas Frederic, Baron Cuvier
The founder of paleontology
Developed the study of fossils by expanding the Systema Naturae to include distinct forms such as dinosaurs
Observed the pattern that fossils and living organisms become increasingly dissimilar as we go back in time
Suggested that the fauna of the past was once dominated by reptiles, not mammals, based on dinosaur bones
Key to the development of the idea that extinction is a widespread process
Directional selection
Removes genetic variation in populations while random mutation replenishes that variation
Null model
A model that predicts the data you should expect to see if a process is not working—meaning, when a particular causative agent is not impacting the situation.
Hardy-Weinberg Principle
A null model stating that if no selection, drift, gene flow, or mutation occur, and if mating is random with respect to the gene in question, then allele frequencies will not change from one generation to the next and genotype frequencies will be in the proportions p2, 2pq, q2, where p and q are the frequencies of two alleles at that gene.
Assortative mating
Mate choice based on similarities or dissimilarities in phenotype.
Inbreeding
Mating among relatives.
Gene pool
A concept used in predicting the results from random matings among all of the individuals in a population, by supposing that all gametes are tossed into "pool" and then drawn out at random to produce offspring.
Genetic bottleneck
The phenomenon in which a population lineage shrinks to a small size for a period, causing that population to lose genetic variation
Genetic drift
Random change in allele frequencies in a population over time
Genetic load
The frequency of deleterious alleles that have accumulated in a population
Beneficial mutation
A new allele that enhances the fitness of organism
Deleterious mutation
A new allele that decreases organismal fitness
Directional selection
Selection that arises when one allele consistently raises fitness; eventually the beneficial allele is expected to become fixed in the population
Mutation
A change in a gene sequence; arises independent of the needs of the organism.
Neutral mutation
A new allele with neither a beneficial nor a deleterious effect
Relative fitness
The fitness of a given genotype divided by the fitness of a reference genotype, which relative fitness is assigned to be 1.0
Carrier
An individual who is heterozygous for a disease
Overdominant selection
Selection in which heterozygote genotypes have the highest fitness
Trans-specific polymorphism
A set of alleles that are shared between closely related species; they arose before speciation and were maintained as polymorphisms
Underdominant selection
Selection in which heterozygote genotypes have the lowest fitness
Continuous traits
Genes not controlled by a single locus, but instead multiple loci, that have a bell shaped curve in which the intermediate trait is usually expressed
Allele frequency
In a population, the proportion of all alleles at a locus that are of a particular type
Hardy-Weinberg Law
The law that, when a few key assumptions hold, makes it is possible to predict the genotype frequencies in a population for the next generation, based on the allele frequencies in the current generation
Locus
A place in the genome where alleles reside; in diploids, each individual has two alleles per locus
Phenotype
The physical and behavioral characteristics of an organism that result from the interaction of the organism’s genotype and its environment
Genotype
Genetic makeup of an organism
Population
A group of interbreeding organisms
Selfing
When an organism reproduces with itself, providing both the egg and sperm components
Allopatric speciation
Speciation driven by geographic isolation
Assortative mating
The phenomenon in which individual organisms tend to mate with other organisms with trait values like theirs
Stabilizing selection
Selection that disfavors extreme trait values and favors trait values towards the center of the trait distribution
Biological Species Concept
The view that species are defined by the ability of their members to reproduce with one another and to be unable to reproduce with the members of other species
Clinal variation
Gradual changes in traits as a function of geographical separation
Discrete variation
Genetic variation among geographically separated populations, where each population contains genetically similar individuals
Extrinsic reproductive isolation
The phenomenon in which two organisms are unable to reproduce due to geographic separation
Intrinsic reproductive isolation
The phenomenon in which two organisms are no longer able to reproduce, even when they are allowed to encounter one another
Phylogenic Species Concept
The view that species are clades (monophyletic groups), like other in the taxonomic hierarchy (genera, subspecies, etc.), that biologists have chosen to assign to the species rank for practical reasons
Speciation
The splitting of an ancestral species into descendant species
Sympatric speciation
Speciation without geographical separation, driven by assortative mating within a population
Disruptive selection
Selection that favors trait values at both extremes of the trait value distribution
Heritability (h²)
The fraction of the variation in a population that can be explained by genetics
Response to selection, r
The amount the mean trait value in a population changes after one generation
Stabilizing selection
Selection that disfavors extreme trait values and favors trait values towards the center of the trait distribution
Standard deviation
Square root of the variance
Variance
A measure of the spread of the distribution of a trait values in a population (technically, the sum of the squared deviations from the mean value)
Altruistic behavior
Actions (or tendencies to act) that result in the organism exhibiting the behavior lowering its own fitness while increasing the fitness of other organisms within its population
Exaggerated secondary sexual selection
Dramatic traits in an organism that lower the organism’s viability but evolve because they are favored by sexual selection
Polygamy
The phenomenon in which one male can mate with many females
Runaway sexual selection
the phenomenon in which secondary sexual characteristics become exaggerated due to feedback between male traits and female preferences (or, more rarely, female traits and male preferences)
Covergent evolution.
The phenomenon observed in phylogenetic trees where the same trait evolves separately in more than one lineage; although they appear similar, these traits are not homologous
Homologous
A descriptor of a character shared in separate species that was inherited from the common ancestor of those species
Principle of parsimony
The idea that given multiple alternative hypotheses, the most likely hypothesis is the one that makes the fewest assumptions; in relation to phylogenies, parsimony favors the scenario that invokes the minimum number of evolutionary changes (character loss/gain)
Reversal
The phenomenon in which an ancestral trait was lost and then re-evolved along a lineage
Separate ancestry
The alternative hypothesis to common ancestry, which proposes that each living taxon has an independent origin
Anoxygenic photosynthesis
The reduction of carbon dioxide to organic molecules using light energy in which something other than water is the electron donor and oxygen gas is not released
Autogenous hypothesis for eukaryotes
The theory that mitochondria and nucleus both evolved within the same eukaryotic lineage
Cyanobacteria
A clade of bacteria characterized by its ability to perform oxygenic photosynthesis
Endosymbiosis
The phenomenon in which a prokaryotic cell comes to live and divide within a host cell
Endosymbiotic theory for eukaryotes
The theory that mitochondria are derived from endosymbiotic bacteria taken up by the host cell whose genome is found in the eukaryotic nucleus
Eukaryotic cells
A category of cells characterized by an outer membrane and many internal membrane-bound compartments including the mitochondria and nucleus
Inside-out model
A theory for development of eukaryotic internal compartments that suggests that the outer plasma membrane of a prokaryotic ancestor was pushed outward and ultimately fused to create the cytoplasm and plasma membrane of eukaryotes.
LUCA (last universal common ancestor)
The most recent ancestor of all known life
Organelle
Membrane-bound compartment of a eukaryotic cell
Outside-in model
A theory for development of eukaryotes that starts with the production of vesicles within the cytoplasm by internalization
Oxygenic photosynthesis
The reduction of carbon dioxide to organic molecules using light energy in which water is the electron donor and oxygen gas is released
Prokaryotic cells
A category of cells characterized by an outer membrane containing no internal membrane structures
Metazoan
a multicellular animal that is made up of cells differentiated into tissues and organs.
Opisthokont clade
a broad group of eukaryotes that includes Animals and Fungi.
Bilateral symmetry
body plan where an organism has a head and a tail, a left and right, and an up and down.
Cambrian Period
a period of rapid diversification of complex animal life when all major bilaterian phyla appeared.
Evolutionary arms race
this occurs when two interacting species develop adaptations and counter-adaptations to each other.
Red Queen Principle
an evolutionary concept where species must continually adapt and evolve to survive, or they risk extinction.
Arthropods
invertebrates with segmented bodies, an exoskeleton, and paired, jointed appendages.
Tetrapods
vertebrates with four limbs.
Amphibians
cold-blooded vertebrates with scale-less, moist skin who spend part of their life in water and part on land.
Amniotes
animals characterized by having an egg with an amnion (a thin membrane that surrounds the fetus within the amniotic sac)
Admixture
The mating of two individuals from genetically distinct group
Diurnal
The characteristic of being active during the day
Introgression
The transfer of genetic information between species