Speciation
Introduction to Speciation
- Speciation: the process in which biological lineages diverge due to reproductive isolation, resulting in new species
- AKA macroevolution
Defining a Species:
- Species Concepts: proposals for how to define a species
- Morphological species concept:
- Defines a species based on observable morphological characteristics
- Ex: body shape, size, color, and other structural features
- Carolus Linnaeus
- Two organisms are a species if they look alike
- Problems:
- Members of the same species may not look alike: sexual dimorphism
- Members of different species may look similar: cryptic species look alike but do not interbreed
- Biological species concept:
- Defines a species as groups of actually or potentially interbreeding natural populations which are reproductively isolated from other such groups
- Ernst Mayr
- Two species are the same species if they can mate with one another
- Reproductive isolation: a situation in which two groups of organisms are capable of exchanging genes
- Problems:
- The definition does not apply to asexual organisms, like prokaryotes
- The definition cannot be applied to extinct species, as fossils do not tell us about sexual isolation of animals
- Many species hybridize, like mules
- Lineage species concept:
- Defines a species as two groups that share a branch on the tree of life
- Relies on molecular similarity and phylogeny
- Advantage: accommodates asexual organisms and hybridization
- Disadvantage: may classify organisms as two different species based on unimportant genetic differences
The Role of Reproductive Isolation:
Reproductive isolation is the most important factor in speciation
Speciation requires interruption in gene flow between two groups
Ways gene flow can be disrupted:
- Behavioral - assortative mating
- Geographical - habitat selection
- Genetic
- Physiological barriers
Dobzhansky-Muller Method: simple model that explains how a single lineage can split into two reproductively isolated species
- Explains role of mutation on speciation
- Steps:
- Something causes two groups to become reproductively isolated
- Over evolutionary time, a different mutation may occur at a different loci in each group
- If these mutations convey an advantage, they will become fixed in the population over time
- The mutant allele from one group may then be incompatible with the mutant allele in the other group, preventing the groups from interbreeding
- Applies to major chromosomal rearrangements and point mutations
The Relationship Between Genetic Divergence and Reproductive Isolation:
- Reproductive isolation tends to increase as species diverge genetically
- Two groups of organisms are more likely to be classified as separate species if their DNA is similar
- Can develop quickly over a course of a single generation or can develop over the course of millions of years
- Genetically distant pairs = high reproductive isolation
- Genetically similar pairs = low reproductive isolation
- Yet, it is still possible to have high levels of reproductive isolation even without much genetic divergence
Incomplete Hybridization and Hybrid Zones:
- Hybrid zones: regions where populations of two closely related species overlap, such that they can hybridize
- Existence of hybrid zones indicates that reproductive isolation between two populations is incomplete
- Usually very narrow, bc strong selective pressure against hybrids
- Natural selection favors mechanisms that prevent species from creating a hybrid in the first place
Allopatric vs. Sympatric Speciation:
- Speciation requires the isolation of gene flow (migration)
- 2 ways that can happen: allopatric or sympatric
Allopatric Speciation:
- Allopatric speciation: occurs when a population is divided by a physical or geographic barrier, forming two reproductively isolated populations that evolve independently
- Major form of speciation in most organisms (especially animals)
- Examples: climate change, glacial advance and retreat, a change in sea levels, continental drift
- Allopatric speciation can also result from founder events: cases in which individuals from one population colonize another region
- Drosophila flies: Hawaii is home to more than 800 species of Drosophila flies, due to around 45 founder events
- Galapagos finches: Finches on each island developed specific characteristics as a result of different environmental conditions from island to island
- Examples of adaptive radiations, which occur when a single species undergoes multiple speciation events so that each newly formed species is able to occupy a specific niche in the environment
Sympatric Speciation:
- Sympatric speciation: occurs when species arise from a population that remains connected
- More common in plants than animals
- 3 major causes of sympatric speciation: disruptive selection, assortative mating, and polyploidy
- Disruptive selection: favors two extreme phenotypes and can result in divergence between two species
- Assortative mating: assortative mating (nonrandom mating) could also result in disruptive selection
- Polyploidy: occurs when a mistake during cell division results in an individual with cells that have more than one copy of the genome (viable = can live and reproduce, but genetically isolated = cannot reproduce with normal diploid individuals)
- In plants, they are able to self fertilize, create offspring and thus propagate a new species
Reproductive Isolation:
- Incipient species: refer to species that are in the process of diverging into separate species, but that can currently interbreed
- When they come back to reproduce (hybridization) two possibilities:
- Reinforcement: hybrids may be less reproductively fit, hybrid offspring may be weak or sterile
- Natural selection will favor parents who mate with their own species over parents who hybridize
- Known as post-zygotic reproductive isolation
- Merger: sometimes, hybrids are no less fit than offspring of non-hybridizing parents
- Populations may interbreed to a point that the formerly separate species become a single species again
- Reproductive barriers:
- Prezygotic isolating mechanisms: prevent hybridization in the first place
- Postzygotic isolating mechanisms: operate after zygote development and reduce the fitness of the hybrid offspring, cause selection against hybridization, leading to reinforcement of prezygotic isolating mechanisms
Prezygotic Isolating Mechanisms:
- Prezygotic isolating mechanisms: reproductive barriers that prevent mating or fertilization between two individuals
- Mechanical isolation: some animals are simply not physically compatible with other animals due to anatomical differences
- Temporal isolation: some organisms are isolated such that they never have the opportunity to reproduce with one another due to timing of reproduction
- Behavioral isolation: some animals have mating rituals that must occur before mating, and pairing of mates does not occur unless the correct mating rituals or signals are performed
- Habitat isolation: some animals are isolated by space such that they never have the opportunity to come in contact with one another
- Gametic isolation: some animals produce sperm and egg cells that will simply not fuse when they come into contact
Postzygotic Isolating Mechanisms:
- Postzygotic isolating mechanisms: reproductive barriers that prevent interbreeding after the development of the zygote and reduce fitness of hybrid offspring
- Low hybrid zygote viability: Sometimes, an offspring is created but is not viable and die soon after fertilization
- Low hybrid adult viability: when the offspring mate with one another or with either parent species, offspring in the next generation become feeble or sterile
- Hybrid infertility: form offspring that can live but not reproduce
Natural Selection and Reproductive Isolation:
- Natural selection favors prezygotic rather than postzygotic
- A lot more efficient to prevent reproduction from occuring in the first place through prezygotic
- Populations living together (sympatry) tend to evolve more effective prezygotic reproductive barriers than populations living apart (allopatry)