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Concept of Speciation
If gene flow stops, allele frequencies in isolated populations can diverge. the populations begin to evolve independently, divergence will occur as a result of mutation, natural selection, and genetic drift, eventually leading to the formation of new species.
Speciation
is a splitting event that creates two or more distinct species from a single ancestral species.
Species
defined as evolutionarily independent population or group of population
3 approaches of identifying species
Biological species concept
morphological species concept
phylogenetic species concept
Biological Species Concept
the main criterion for identifying species is reproductive isolation which means
results in lack of gene flow between populations
members in population do not interbreed, or fail to produce viable, fertile offspring after mating.
mechanisms that stop gene flow between population
are known as reproductive isolating mechanisms, which can be prezygotic or postzygotic isolation.
Prezygotic isolation
individuals or different species are prevented from mating successfully due to various barriers such as temporal, habitat, behavioral, mechanical, or gametic barrier.
Postzygotic isolation
the hybrid offspring do not survive or reproduce successfully due to various mechanisms such as hybrid viability, hybrid sterility.
Disadvantages of Biological Species concept
reproductive isolation cannot be evaluated in fossils like Trilobites and species that preproduce asexually
Temporal Isolation
population are isolated because they breed at different times ex: Spotted skunks they mate different time of the year
Habitat isolation
population are isolated because they breed in different habitats ex: garter snakes, different species live in different habitats
behavioral isolation
populations do not interbreed because of different mating behaviors or rituals, such as specific calls or dances that attract mates. ex: Blue and red footed booby - different courtship
Gametic Isolation
mating fails because sperm and egg are incompatible, preventing fertilization. This can occur in many species, including some types of sea urchins.
Mechanical Isolation
mating fails because male and female reproductive structures are incompatible ex: in drosophila species there are subtle differences in genitalia that prevent successful mating.
Hybrid (in) viability
hybrid offspring do not develop normally and die as embryos ex: salamanders
Hybrid Sterility
hybrid offspring mature but are sterile as adults ex: mules, which are hybrids of horses and donkeys.
Haldane’s Rule
when in F1 offspring of two different animal races one sex is absent, rare, or sterile that sex is the heterozygous sex
males are sterile in XY taxa (flies, mammals)
females are sterile in ZW taxa (birds, butterflies)
Morphospecies Concept
individual lineages differ in size, shape, or other morphological features
it is widely applicable
when there is no data on the extent of gene flow
Equally applicable to sexual, asexual, and fossil species
Disadvantages of Morphospecies concept
one polymorphic species maybe classified as more than one species
it cannot identify cryptic species that differ in non morphological traits
The features used to distinguish species under this concept is subjective
Phylogenetic species Concept
identifies species based on evolutionary history
based on the rationale that all species are related by common ancestry
Only on phylogenetic trees, a monophyletic group consists of an ancestral species and all its descendants.
Also called a clade or lineage
monophyletic group
defined as an ancestral population and all descendants
synapomorphy
defined as a trait unique to a monophyletic group (unique forms)
Advantages of phylogenetic species concept
it can be applied to any type of population (e.g fossils, sexual or asexual)
It is logical because different species have different synapomorphies due to lack of gene flow and independent evolution
Disadvantages of phylogenetic species concept
phylogenies are currently available for only a tiny (through growing) subset of population on the tree of life
critics point out that it would probably lead to recognition of many more species than either of the other species concepts.
Why do researchers use all 3 species concepts?
crosses to test compatibility
population genetics to characterize genetic exchange
phylogenetic to identify evolutionary history
3 modes of speciation
allopatric speciation
parapatric speciation
sympatric speciation
Allopatric speciation
speciation that begins with geographic isolation
It occurs when a populations become geographically separated.
population that live in different areas are in allopatry
the most common mode
geographic isolation occurs in 2 different ways
dispersal
vicariance
Dispersal
A population moves to a new habitat, colonizes it, and found a new population
Vicariance
a physical barrier splits a population into subgroups that are physically isolated from each other.
Biogeography
the study of how species and populations are distributed geographically, it can tell us how dispersal and vicariance events occur
Allopatric Speciation by Dispersal
Colonization events can lead to speciation
Colonists represented a new allopatric population, but could have led to speciation
The Grants concluded that both mechanisms contributes the change in beak size in the large ground finches
New population is not a sepatate species yet, because there is still some gene flow
Over time the populations could continue to diverge
Allopatric Speciation by vicariance
• When there is a physical barrier such as a mountain range uplifting or a river splitting the geographic range of a species, vicariance has occurred.
Example: Researchers compared the DNA sequences of trumpeters from different areas of the Amazon basin to the geological events that occurred
• They found evidence of isolation of populations by vicariance
• Initially, the formation of the Amazon split the ancestral population
• The formation of the river systems then subdivided the population
Parapatric Speciation
Genetic isolation happens when populations change gradually along a cline
Geographic features such as latitude, or elavation impose a gradient of selection pressures
Phenotypic and genetic change result
cline
spatial gradient of character change along an environmental variable
sympatry
Populations or species that live in the same geographic area - close enough to interbreed
Isolation and Divergence in sympatry
researchers believed that speciation could not occur among sympatric populations because gene flow would overwhelm any differences among populations,
Sympatric Speciation has two types of events
External
Disruptive selection
Internal
chromosomal mutations
Disruptive Selection
different ecological niches or mate preferences
niche
the range of ecological resources that a species can use and the range of conditions it can tolerate.
Sympatric speciation by polyploidization
Polyploidy can cause speciation where an error in meosis or mitosis result in more than two sets of chromosomes
Autopolyploid
a doubling of chromosome number
Chromosomes are all from the same species
New tetraploid species may be created when two diploid species hybridize.
Offspring are usually sterile. Offspring has two copies of each of two sets of chromosomes.
Allopolyploid
parents of different species mate and an error in mitosis occurs, resulting in viable non sterile offspring
Produces offspring with two different sets of chromosomes.
tetraploid
individuals can sucessfully breed eith other tetraploids but not with diploids
Advantages of polyploid
have higher levels of heterozygosity
can tolerate higher levels of self-fertilization because they are not as affected by inbreeding depression
genes on duplicated chromosomes cab diverge independently
What happens when isolated populations come into contact?
If separation has occurred and prezygotic isolation exists, then mating between the population is rare, and populations continue to diverge
What happens when isolated populations do not come into contact?
When prezygotic barriers do not exist, populations may interbreed and potentially erase distinctions between the two populations. Other possible outcomes are reinforcement, development of hybrid zones.
Reinforcement
Natural selection of traits that prevent interbreeding among populations
selection for prezygotic isolating mechanism
evidence of reinforcement
Several studies show that prezygotic isolation mechanisms are found between species that overlap geographically.
Hybrid Zones
A Geographic area where interbreeding between two populations occurs, which sometimes shows that the hybrid offspring of separated populations can mate and produce viable, fertile offspring. Their hybrid zones can be either narrow or wide, long or short-lived
stable in one place or move over time
Hybrid zones data shows
Data from the mitochondrial DNA show that most hybrids form when Townsend’s warbler males mate with hermit warbler females. The males invade hermit territories, drive off hermit males, and mate with hermit females.
Phylogeny
The evolutionary history of a group of organisms
Phylogenetic Tree
is a graphical summary of history, it can show evolutionary relationship among genes, species, higher taxa
Tree of life
can be used in taxonomy to define species
can be used in medicine to study the spread of disease
can aid in identifying species that are in conservation priority
can be used in agriculture to identify wild relatives for breeding with current crops
A branch in the phylogenetic tree represents
a population through time
A node in the phylogenetic tree represents
A point where a branch splits - that is hypothetical most recent ancestor, represent speciation events
a tip in the phylogenetic tree represents
The endpoint of a branch which is a living or extinct taxon
A root in the phylogenetic tree represents
ancestral branch
An out group in the phylogenetic tree represents
A taxon that is separated before the taxa of interest did,when they evolve they donot represent other taxa in the tree
Polytomy
is a node that divides into 3 or more branches
Taxa
Always located in the brach tips, never within the tree because none of the taxa are presumptive ancestors of others. Related taxa are sister groups that share a common ancestor.
How do biologists estimate phylogenies?
character-state data
genetic distance data
Character state data
A character or trait is any genetic, morphological, physiological or behavioral characteristic to be studied
ancestral trait
character that existed in an ancestor
derived trait
One that is a modified form of an ancestral trait, found in a descendant species.
homology
similarity in traits due to common ancestry
Synapomorphy
a shared, derived trait
homoplasy
Similarity in organisms due to reasons other than common ancestry
polyphyletic group
An unnatural group that does not include the most recent common ancestor
paraphyletic group
A group that includes a common ancestor and some, but not all, of its descendants.
homologies
traits shared among related species
homoplasies
traits shared among unrelated species
convergent evolution
Similar design solutions to environmental challenges that arise independently in different evolutionary lineages.
Parsimony
The principle of logic states that the most likely explanation or pattern is the one that implies the least number of changes. the trees with the least number of evolutionary changes is likely the one that most accurately reflects what occured during evolution
Principle of parsimony
in the absence of other data, the best hypothesis minimizes the number of evolutionary changes (minimizes the number of extra changes on tree due to homoplasy)
molecular clock
a technique used by evolutionary biologists to estimate the timing of evolutionary events by measuring the accumulation of genetic mutations in DNA or protein sequences
Fossil Record
It is the total collection of fossils that have been found throughout the world
Fossil
physical evidence from an organism that lived in the past
The Fossil record provides evidence about
what organisms that lived in the past looked like
where they lived
when they existed
how do fossils form?
most of the processes that form fossils begin when part or all of an organism is buried by sediment, which then hardens over time, preserving the organism's remains.
intact fossils
Decomposition does not occur, and organic remains are preserved
compression fossils
when sediments accumulate on top of the material and compress it into a thin carbonaceous film
Cast fossils
form when the remains decompose after burial and dissilved minerals create a cast in the remaining hole
per mineralized fossils
when the remains decompose slowly and dissolved minerals slowly infiltrate the cells’ interiors and harden into stone
Trace Fossils
When sedimentation and mineralization preserve indirect evidence of an organism
how do fossils form?
fossils form only under ideal conditions
they must be buried rapidly
they must decompose slowly
limitations of fossil record
habitat bais
taxonomic and tissue bias
temporal bias
abundance bais
habitat bais
Organisms that live where sediment is actively being deposited are more likely to fossilize than those living aboveground. in these habitats burrowing organisms are more likely to fossilize compared to organisms living aboveground
taxonomic and tissue bais
some organisms those with hard shells such as bones or shells are more likely to decay slowly and leave fossil evidence
temporal bias
more recent fossils are more common than ancient fossils
Abundance bais
Organisms that are abundant, widespread, and present for a long time leave evidence much more often than do species that are rare, local, or ephemeral
precambrian era
interval from earth’s formation to the apperance of most animal groups about 543 million years ago
unicellular until late
phanerozoic eron
Spans the interval between ~543 mya and the present
Is divided into three eras—the Paleozoic, the Mesozoic,
and the Cenozoic
Paleozoic Era
Begins with the appearance of most major animal lineages
• Includes the initial diversification of animals, land plants, and fungi
• Land animals first appear
• Ends with the obliteration of almost all multicellular life-forms at
the end of the Permian period
• ~543 mya – 248 mya
Mesozoic era
Begins with the extinction events at the end of the Paleozoic and
ends with the extinction of dinosaurs
• Dinosaurs and gymnosperms were the most dominant terrestrial
vertebrates and plants, respectively
• 252 mya – 66 mya
cenozoic era
mammals and angiosperms were the largest terrestrial vertevrates and plants, respectively
Adaptive Radiation
the rapid production, from a single lineage, of many descendant species
Extinctions
dramatic reduction in the number of species due to environmental change
Two general mechanisms can trigger adaptive radiations
Extrinsic factors such as favorable new conditions in the
environment (Ecological opportunity)
Intrinsic factors such as evolution of key morphological,
physiological, or behavioral traits (Key innovation)
Ecological opportunity
the availability of more or new types of resources—has driven a wide array of adaptive radiations
By being able to disperse into new environments, they could radiate into lots of species