Speciation

What Is a Species?
One of the best definitions is that of the evolutionary biologist
Ernst Mayr:
A species is an actually or potentially interbreeding
population that does not interbreed with other such
populations when there is opportunity to do so.
Note: sometimes breeding may take place (as it can between a horse and a donkey) but
if so, the offspring are not so fertile and/or well adapted as the parents (the mule
produced is sterile).

Allopatric Speciation: the Role of Isolation in

Speciation
The formation of two or more species often (some workers think always!) requires
geographical isolation of subpopulations of the species. Only then can natural selection
or perhaps genetic drift produce distinctive gene pools.

Link to discussion of gene

pools and the forces that alter
them.

It is no accident that the various races (or "subspecies") of animals almost never occupy
the same territory. Their distribution is allopatric ("other country").

The seven distinct subspecies or races of the yellowthroat Geothlypis trichas in the
continental U.S. would soon merge into a single homogeneous population if they
occupied the same territory and bred with one another.

Darwin's Finches
As a young man of 26, Charles Darwin visited the Galapagos Islands off the coast of
Ecuador.
Map

Among the animals he studied were 13 species of finches found nowhere else on earth.

 Some have stout beaks for eating seeds of one size or another.
 Others have beaks adapted for eating insects.
 One (#7) has a beak like a woodpecker's. It uses it to drill holes in wood, but
lacking the long tongue of a true woodpecker, it uses a cactus spine held in its
beak to dig the insect out.
 One (#12) looks more like a warbler than a finch, but its eggs, nest, and
courtship behavior is like that of the other finches.
Darwin's finches. The finches
numbered 1–7 are ground finches.
They seek their food on the ground
or in low shrubs. Those numbered
8–13 are tree finches. They live
primarily on insects.

1. Large cactus finch (Geospiza

conirostris)
2. Large ground finch (G.
magnirostris)
3. Medium ground finch (Geospiza
fortis)
4. Cactus finch (G. scandens)
5. Sharp-beaked ground finch (G.
difficilis)
6. Small ground finch (G.
fuliginosa)
7. Woodpecker finch (Cactospiza
pallida)
8. Vegetarian tree finch (Platyspiza
crassirostris)
9. Medium tree finch
(Camarhynchus pauper)
10. Large tree finch
(Camarhynchus psittacula)
11. Small tree finch (C. parvulus)
12. Warbler finch (Certhidia
olivacea)
13. Mangrove finch (Cactospiza
heliobates)

(From BSCS, Biological Science:

Molecules to Man, Houghton
Mifflin Co., 1963)
Since Darwin's time, these birds have provided a case study of how a single species
reaching the Galapagos from Central or South America could — over a few million
years — give rise to the 13 species that live there today.

Several factors have been identified that may contribute to speciation.

Ecological opportunity
When the ancestor of Darwin's finches reached the Galapagos, it found
 no predators (There were no mammals and few reptiles on the islands.)
 few, if any, competitors. There were only a handful of other types of songbirds.
In fact, if true warblers or woodpeckers had been present, their efficiency at
exploiting their niches would surely have prevented the evolution of warblerlike
and woodpeckerlike finches.
Geographical Isolation (allopatry)
The proximity of the various islands has permitted enough migration of Darwin's
finches between them to enable distinct island populations to arise. But the distances
between the islands is great enough to limit interbreeding between populations on
different islands. This has made possible the formation of distinctive subspecies (=
races) on the various islands.

The importance of geographical isolation is illuminated by a single, fourteenth, species

of Darwin's finches that lives on Cocos Island, some 500 miles to the northeast of the
Galapagos.

The first immigrants there must also have found relaxed selection pressures with few
predators or competitors.

How different the outcome, though. Where one immigrant species gave rise to at least
13 on the scattered Galapagos Islands, no such divergence has occurred on the single,
isolated Cocos Island.

Evolutionary Change
In isolation, changes in the gene pool can occur through some combination of
 natural selection
 genetic drift
 founder effect

These factors may produce distinct subpopulations on the different islands. So long as
they remain separate (allopatric) we consider them races or subspecies. In fact, they
might not be able to interbreed with other races but so long as we don't know, we
assume that they could.

How much genetic change is needed to create a new species?

Perhaps not as much as you might think. For example, changes at one or just a few gene
loci might do the trick. For example, a single mutation altering flower color or petal
shape could immediately prevent cross-pollination between the new and the parental
types (a form of prezygotic isolating mechanism).

Reunion
The question of their status — subspecies or true species — is resolved if they ever do
come to occupy the same territory again (become sympatric). If successful
interbreeding occurs, the differences will gradually disappear, and a single population
will be formed again. Speciation will not have occurred.

If, on the other hand, two subspecies reunite but fail to resume breeding, speciation has
occurred and they have become separate species.
 An example: The medium tree finch Camarhynchus pauper is
found only on Floreana Island. Its close relative, the large tree
finch, Camarhynchus psittacula, is found on all the central islands
including Floreana.

Were it not for its presence on Floreana, both forms would be

considered subspecies of the same species. Because they do coexist
and maintain their separate identity on Floreana, we know that
speciation has occurred.

Isolating Mechanisms
What might keep two subpopulations from interbreeding when
reunited geographically? There are several mechanisms.

Prezygotic Isolating Mechanisms

These act before fertilization occurs.

 Failure to elicit mating behavior. On Floreana, C. psittacula has a longer beak
than C. pauper, and the Grants have demonstrated that beak size is an important
criterion by which Darwin's finches choose their mates.
 Two subpopulations may occupy different habitats in the same area and thus fail
to meet at breeding time.
 In plants, a shift in the time of flowering can prevent pollination between the
two subpopulations.
 Structural differences in the sex organs may become an isolating mechanism.
 The sperm may fail to reach or fuse with the egg.

Postzygotic Isolating Mechanisms

These act even if fertilization does occur.

Even if a zygote is formed, genetic differences may have become so great that the
resulting hybrids are less viable or less fertile than the parental types. The sterile mule
produced by mating a horse with a donkey is an example.

When Drosophila melanogaster attempts to mate with its relative Drosophila

simulans, no viable males are produced. Mutations in a single gene (encoding a
component of the nuclear pore complex) are responsible.

Speciation by Hybridization
Although hybridization between related species of angiosperm also causes reduced
fertility, it is sometimes followed by a doubling of the chromosome number. The
resulting polyploids are now fully fertile with each other although unable to breed with
either parental type. A new species has been created. This appears to have been a
frequent mechanism of speciation in angiosperms.
Link to a discussion of polyploidy and an example of its role in speciation.

Even without forming a polyploid, interspecific hybridization

can occasionally lead to a new species of angiosperm. Two
species of sunflower, the "common sunflower", Helianthus
annuus, and the "prairie sunflower", H. petiolaris, grow widely
over the western half of the United States. They can interbreed, but only rarely are
fertile offspring produced.

However, Rieseberg and colleagues have found evidence that successful hybridization
between them has happened naturally in the past. They have shown that three other
species of sunflower (each growing in a habitat too harsh for either parental type) are
each the product of an ancient hybridization between Helianthus annuus and H.
petiolaris. Although each of these species has the same diploid number of
chromosomes as the parents (2n = 34), they each have a pattern of chromosome
segments that have been derived, by genetic recombination, from both the parental
genomes. They demonstrated this in several ways, notably by combining RFLP analysis
with the polymerase chain reaction (PCR).

They even went on to produce (at a low frequency) annuus x petiolaris hybrids in the
greenhouse that mimicked the phenotypes and genotypes of the natural hybrids. (You
can read about the results of these monumental studies in the 29 August 2003 issue of
Science.)

So, at least in angiosperms, speciation can occur as a result of hybridization between

two related species, if the hybrid
 receives a genome that enables it to breed with other such hybrids (these
angiosperms can fertilize themselves) but
 not breed with either parental species;
 can escape to a habitat where it does not have to compete with either parent,
 is adapted to live under those new conditions.

Intense Competition
The process of speciation is often hastened when two formerly isolated groups are
reunited. Even if they no longer interbreed, they are probably still similar in many ways,
including their requirements for the necessities of life.

Thus the reuniting of the two groups may create an intense selection pressure leading to
one of two possible outcomes.

1. The competition may be so intense that one species becomes eliminated entirely;
that is, it is driven to extinction on that island. This may have occurred to C.
pauper on some of the central islands.
2. Alternatively, the increased selection pressure may lead to character
displacement and so lessen the competition between them.
 The range of beak sizes of C. pauper on Floreana and
C. psittacula on Isabela is roughly the same. This
reflects the fact that the two species feed on the same type
and size of insect.

On Floreana, however, character displacement has

occurred: C. psittacula has a substantially larger beak than C. pauper does, and
thus differs enough in its food requirements for the two species to coexist there.

The histogram shows the data. The larger beak of C. psittacula on Floreana
reduces the competition with C. pauper (which is not found on Isabela).
(Redrawn with permission from David Lack, Darwin's Finches, Cambridge
Univ. Press, 1947. The number of collected specimens (16) of C. psittacula on
Floreana was too small to compute a reliable percentage but all fell within the
range indicated.)

Adaptive Radiation
The process described above can occur over and over. In the case of Darwin's finches, it
must have been repeated a number of times forming new species that gradually divided
the available habitats between them. From the first arrival have come a variety of
ground-feeding and tree-feeding finches as well as the warblerlike finch and the tool-
using woodpeckerlike finch.

The formation of a number of diverse species from a single ancestral one is called an
adaptive radiation.

House mice on the island of Madeira

A recent (13 January 2000) report in Nature describes a study of house mouse
populations on the island of Madeira off the Northwest coast of Africa. These workers
(Janice Britton-Davidian et al) examined the karyotypes of 143 house mice (Mus
musculus domesticus) from various locations along the coast of this mountainous
island. Their findings:
 There are 6 distinct populations (shown by different colors)
 Each of these has a distinct karyotype, with a diploid number less than the
"normal" (2N=40).
 The reduction in chromosome number has occurred through Robertsonian
fusions. Mouse chromosomes tend to be acrocentric; that is, the centromere
connects one long and one very short arm. Acrocentric chromosomes are at risk
of translocations that fuse the long arms of two different chromosomes with the
loss of the short arms.
 The different populations are allopatric; isolated in different valleys leading
down to the sea.
 The distinct and uniform karyotype found in each population probably arose
from genetic drift rather than natural selection.
 The 6 different populations are technically described as races because there is
no opportunity for them to attempt interbreeding.
  However, they surely meet the definition of true species. While hybrids would
form easily (no prezygotic isolating mechanisms), these would probably be
infertile as proper synapsis and segregation of such different chromosomes
would be difficult when the hybrids attempted to form gametes by meiosis.

Sympatric Speciation
Sympatric speciation refers to the formation of two or more descendant species from a
single ancestral species all occupying the same geographic location.

Some evolutionary biologists don't believe that it ever occurs. They feel that
interbreeding would soon eliminate any genetic differences that might appear.

But there is some compelling (albeit indirect) evidence that sympatric speciation can
occur.

The three-spined sticklebacks, freshwater fishes, that have been studied by Dolph
Schluter (who received his Ph.D. for his work on Darwin's finches with Peter Grant)
and his current colleagues in British Columbia, provide an intriguing example that is
best explained by sympatric speciation.

They have found:

 Two different species of three-spined sticklebacks in each of five different lakes.
o a large benthic species with a large mouth that feeds on large prey in the
littoral zone
o a smaller limnetic species — with a smaller mouth — that feeds on the
small plankton in open water.
 DNA analysis indicates that each lake was colonized independently, presumably
by a marine ancestor, after the last ice age.
 DNA analysis also shows that the two species in each lake are more closely
related to each other than they are to any of the species in the other lakes.
 Nevertheless, the two species in each lake are reproductively isolated; neither
mates with the other.
 However, aquarium tests showed that
o the benthic species from one lake will spawn with the benthic species
from the other lakes and
o likewise the limnetic species from the different lakes will spawn with
each other.
o These benthic and limnetic species even display their mating preferences
when presented with sticklebacks from Japanese lakes; that is, a
Canadian benthic prefers a Japanese benthic over its close limnetic
cousin from its own lake.
 Their conclusion: in each lake, what began as a single population faced such
competition for limited resources that
o disruptive selection — competition favoring fishes at either extreme of
body size and mouth size over those nearer the mean — coupled with
o assortative mating — each size preferred mates like it
 favored a divergence into two subpopulations exploiting different food in
different parts of the lake.

 The fact that this pattern of speciation occurred the same way on three separate
occasions suggests strongly that ecological factors in a sympatric population
can cause speciation.

Sympatric speciation driven by ecological factors may also account for the
extraordinary diversity of crustaceans living in the depths of Siberia's Lake Baikal.

Parapatric Speciation
There is another possible way for new species to arise in the absence of geographical
barriers.

 If a population ranges of a vast area (e.g., the Amazon basin in south America)
and
 if the individuals in that population can disperse over only a small portion of this
range,

then gene flow across these great distances would be reduced.

Genetic isolation arising simply from the great distance separating subpopulations could
thus lead to "parapatric" speciation. Some workers feel that the enormous species
diversity of the rain forests of South America (greater than that of Africa and Asia
combined!) arose from parapatric speciation.

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