Chapter 24 Origin of Species

Speciation
Darwin referred to the first

appearance of a new organism

on Earth as the mystery of mysteries. Today, the process by which one species splits into two or more species is called speciation.

Speciation forms a conceptual bridge between microevolution , changes over time in allele frequencies in a population, and macroevolution, the broad pattern of evolution over time.

Biological Species Concept

Simple definition = a species is one kind of organism As described by Ernst Mayr in 1942, species is a group of populations whose members have the potential to interbreed in nature, and produce viable, fertile offspring but do not produce viable, fertile offspring with members of other such groups.

The formation of a new species hinges on reproductive isolation

Reproductive Isolation is barriers that prevent members of two species from producing viable, fertile offspring.

Reproductive Isolation
Barriers block gene flow between the species and limit the formation of hybrids, offspring that result from interspecific mating

What holds the gene pool of a species together even when they are divided up into separate populations? What if the populations are long distances apart? The answer is gene flow, the transfer of alleles between populations.

If researcher, Scott Edwards found one population had an allele that shared a recent common ancestor with alleles found in a different population, he reasoned that gene flow must have occurred.

Do the data indicate gene flow occurred as the transfer of an allele from population A to B or the reverse?

Reproductive Barriers
Prezygotic barriers = block fertilization = no zygote

high step courtship display Mate in winter vs summer Water vs terrestrial

Genitals not well matched

Reproductive Barriers
Postzygotic barriers = after fertilization occurs and the zygote is formed.

Sperm cannot fertilize egg

Incomplete development of hybrids

Viable & fertile hybrids produce small, sterile offspring in next generation

Donkey + horse = sterile mule

Limitations of the Biological Species Concept

Can not be applied to Fossils Can not be applied to organisms that reproduce asexually

Species = populations whose members have the potential to interbreed in nature, and produce viable, fertile offspring

Other Definitions of Species

Morphological Species concept = species have common body shape and other structural features Actually the way most species are distinguished Ecological Species concept = views a species in terms of its ecological niche, the sum of how members of the species interact with the nonliving and living parts of their environment Phylogenetic Species concept = defines a species as the smallest group of individuals that share a common ancestor, forming one branch on the tree of life. More than 20 other species definitions have been proposed

Speciation can occur in two main ways; Allopatric (other country) Speciation Sympatric (same country) Speciation

Allopatric Speciation
Allopatric Speciation occurs when gene flow is interrupted when a population is divided into geographically isolated subpopulations Once geographic isolation has occurred, the separated gene pools may diverge through normal mechanisms; Different mutations arise Natural selection acts on the separated organisms Genetic drift alters allele frequency

Allopatric Speciation
Geographic isolation leads to reproductive isolation. Reproductive isolation may arise as a byproduct of selection or drift having caused the populations to diverge genetically.

This isolation must result from barriers that would prevent reproduction even if the different populations come in contact with each other.

Allopatric speciation of antelope squirrels on opposite rims of the Grand Canyon.

Harriss antelope squirrel inhabits the south rim and white-tailed antelope squirrels live on the north rim. Would birds that live in this area be likely to diverge into new species?

After 40 generations natural selection resulted in divergent evolution Flies raised on starch digested it better and flies raised on maltose digested maltose better

Sympatric Speciation
Sympatric Speciation occurs in populations that live in the same geographic area. Gene flow can be reduced by; Polyploidy Habitat differentiation Sexual selection

Polyploidy
Polyploidy is a condition of having extra sets of chromosomes due to a mistake during cell division.

Second type of polyploidy can occur when two different species interbreed and produce hybrid offspring. Usually sterile because chromosomes cant pair during meiosis. Sometimes a fertile polyploid may develop and since these polyploids are only fertile when mating with each other, they represent a new species.

Polyploid speciation rarely occurs in animals, but it is common in plants. Estimates are that about 80% of Earths plant species are the result of polyploid speciation.

Sexual selection
Sympatric speciation may be driven by sexual selection. How did so many different cichlid fish (up to 600 species) originate in Lake Victoria in East Africa over a period of only about 100,000 years?

Does sexual selection in cichlids result in reproductive isolation? Under normal light female cichlids strongly preferred males of their own species. Under monochromatic orange light the two species are very similar in color. Under orange light females showed no preference in mating. The resulting hybrids were viable and fertile

Hybrid zones
Hybrid zones are regions where members of different species meet and mate, producing at least some offspring of mixed ancestry. Some hybrid zones are narrow bands, like between firebellied and yellow-bellied toad ranges.

Yellow-bellied toad alleles

Three possible outcomes for hybrid zone over time

1) Reinforcement: Strengthening Reproductive Barriers

Example: Two species of European flycatcher In sympatric populations (potential for hybrid zone) females are super good at telling males of each species apart In allopatric populations ??

2) Fusion: Weakening Reproductive Barriers

Fusion occurs when reproductive barriers are so weak that gene flow is common between gene pools and eventually may lead to a new species forming. May be happening in Lake Victoria, where mate choice has kept many of the similar species of cichlids reproductively isolated.

3) Stability: Continued Formation of Hybrid Individuals

Remember the Bombina toads, since the hybrids were at a disadvantage why hasnt reinforcement occurred, with strengthened barriers? In over 20 years of study no evidence of reinforcement. Perhaps the narrowness allows more gene flow from outside the zone, overwhelming selection for isolation within the zone.

Speciation can occur rapidly or slowly and can result from changes in few or many genes.

Time between speciation events = time from formation of a new species until it begins to diverge into more than one species, plus the time it takes for speciation to be complete once divergence has begun. = Time elapsed between New species #1 created until New species #2 created Survey of 84 plants and animals showed the interval ranged from 4,000 to 40 million years. The average was 6.5 million and rarely took less than 500,000 years.

Genetics of Speciation
Evolution of as little as one gene can cause reproductive isolation. In some Japanese snails a single gene controls the direction the shells spiral. When their shells spiral in opposite directions the snails genitals are oriented in a manner that prevents mating.

Studies have shown that a very few genes or many genes can influence the evolution of reproductive isolation and the emergence of a new species.

Bornean clouded leopard