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AIM: How does large evolution and speciation happen?

Evolution creates (and destroys) new species, but What is a species? Its not as straightforward a quetion as most believe.

These are members of different species - eastern (left) and western (right) meadowlark.

What is a Species?

There is only one extant (existing) human species.

What is a Species?

And these are all members of a single species.

Determining What Is and What Isnt a Distinct Species Can Have Economic Consequences

Northern spotted owl (left) and barred owl (right).

What is a Species?
The definition well use is this: A species is a group of individuals capable of interbreeding to produce fertile offspring.

This is the biological species concept. Like all attempts to define a species, it has many problems.

One Problem in the Biological Species Concept

For asexually-reproducing organisms, like these bacteria, what constitutes a species?

and another problem with the Biological Species Concept

Where do you draw the line between different species?

How Many Species Are There?

We dont know. About 2 million species have been described. Estimates of existing species number range from 4 million to 100 million (with 10-15 million being a more commonly considered upper estimate).

How did this diversity of life come to be?

Species = basic unit

Continuous lineage - information passed through genes

Speciation - rise of new species

Two Patterns of Speciation

How Do Species Arise?

The key to speciation is reproductive isolation of populations. There are extrinsic and intrinsic reproductive isolating mechanisms.

Geographic isolation is the primary extrinsic reproductive isolating mechanism.

Process of speciation:
Parent species 2 separate species

I. Barrier

II. Diverge

III. Dont interbreed

Reproductive Isolation May Occur With or Without Geographic Isolation

Allopatric speciation occurs when geographic isolation creates a reproductive barrier (an extrinsic mechanism). Sympatric speciation occurs when a reproductive barrier is created by something other than geographic isolation (intrinsic mechanisms).

Allopatric Speciation
Harris antelope squirrel

White-tailed antelope squirrel

Two species of ground squirrel are postulated to have descended from a common ancestral population that was separated by formation of the Grand Canyon.

Intrinsic Reproductive Isolating Mechanisms Are Always Required for Speciation

Harris antelope squirrel

White-tailed antelope squirrel

Intrinsic mechanisms involve changes to organisms that prevent interbreeding.

In allopatric speciation, intrinsic mechanisms come into play once populations are physically separated.

In sympatric speciation, intrinsic mechanisms are the only ones involved.

Many Intrinsic Reproductive Isolating Mechanisms Drive Speciation

(different habits within an overlapping range)

Many Intrinsic Reproductive Isolating Mechanisms Drive Speciation

Courtship rituals, like these, are critical for mating within a species, but ineffective for attracting members of other species.

Behavioral Isolation Mechanisms

Courtship rituals, like these, are critical for mating within a species, but ineffective for attracting members of other species.

Many Intrinsic Reproductive Isolating Mechanisms Drive Speciation

Many Intrinsic Reproductive Isolating Mechanisms Drive Speciation

Hybrid Infertility Was the Impetus for Cloning a Mule

Speciation Occurs at Widely Differing Rates

A slow rate of speciation evidenced by a living horseshoe crab (13 extant species) and a 300 million year-old fossil species A rapid rate of speciation evidenced by Galapagos finches which have diversified into 13 species within the last 100,000 years.

Speciation Rates

Generalists, like the horseshoe crab, tend to remain as stable species.

Specialists, like the Galapagos finch, tend to be unstable as species. Speciation also becomes rapid when, as occurred with Galapagos finches, new niches become available.

Speciation Dynamics - Gradualism or Punctuated Equilibrium?

Punctuated equilibrium appears to be a more accurate view of speciation dynamics.

Does Evolution Create the Perfect Organism?

No, only better organisms as evolution is constrained by history and buffeted by random events. Essentially, every organism on earth is in significant part a sum of accidents.

Species Come and Go


Best estimates from the fossil record indicate that greater than 99% of species that have exited are now extinct.

A typical lifetime for a species is about 1 million years.

Mass Extinctions Are a Fact of Life

90% of all species that have ever existed are now extinct

The Cretaceous/Tertiary Mass Extinction

Gary Larson

Are We Now Causing a Mass Extinction?

Classifying Lifes Rich Diversity

Why bother? An intrinsic reason is that modern classification systems tell whos related to whom and how we all came to be.

Classifying Lifes Rich Diversity

Why bother?

A practical reason is that if we want to preserve an environment compatible with human life, wed better know whats out there.

The Linnaean Hierarchical Classification System

How Do We Classify Organisms?


Ideally, classification is based on establishing the evolutionary relationships between organisms. The evolutionary relationship between organisms is their phylogeny. Cladistics is the method of classification based on establishing phylogenies (i.e. getting at evolutionary relationships.

Cladistics proceeds by comparing shared ancestral and shared derived characters between sets of organisms.

Cladistics

A phylogeny (cladogram) for vertebrates.

each node indicates a common ancestor

The greater the number of derived characters shared by a pair of organisms, the closer their degree of relationship. The closer the degree of relationship, the closer the most recent common ancestor.

Its Critical (and often difficult) To Distinguish Homology from Analogy

Homologous structures, like the bat wing and gorilla arm, are similar because they are derived by modification of a shared ancestral structure. Homology is the key to establishing phylogenies.

Another Set of Analogies Created by Convergent Evolution

Ocotillo of the US southwest

Allauidia of Madagascar

Results of Cladistic Analyses Sometimes Run Counter to Classical Classification Schemes


Which pair is more closely related? A lizard/crocodile or bird/crocodile?

Cladistic analysis indicates that the bird/crocodile pair is more closely related.

DNA Hybridization:
uses DNA similarity between species

Which species are the closest living relatives of modern humans?


Gorillas Humans

Chimpanzees
Bonobos Orangutans Humans
15-30 MYA 0 14 MYA

Chimpanzees
Bonobos

Gorillas
Orangutans
0

The pre-molecular view was that the great apes (chimpanzees, gorillas and orangutans) formed a clade separate from humans, and that humans diverged from the apes at least 15-30 MYA.

Mitochondrial DNA, most nuclear DNA-encoded genes, and DNA/DNA hybridization all show that bonobos and chimpanzees are related more closely to humans than either are to gorillas.