No.

6
Patterns of Descent with Modification
Attempts to define the concept of species date back to the Greek philosophers Plato and Aristotle, who viewed the world as we know it as a
flawed shadow of the eternal and immutable world of ideas. Indeed, the word “species” originates from the Latin “kinds” which is a
translation of the Greek word eidos (idea). Ernst Mayr played a central role in the establishment of the general concept of species as
metapopulation lineages, and he is the author of one of the most popular of the numerous alternative definitions of the species category.
According to him, “Species are groups of interbreeding natural populations that are reproductively isolated from other such groups.” Another
definition based on George Gaylord Simpson, “species is a lineage (an ancestral-descendant sequence of populations) evolving separately
from others and with its own unitary evolutionary role and tendencies.” Similarly, according to Leigh Van Valen, “a species is a lineage (or a
closely related set of lineages) which occupies an adaptive zone minimally different from that of any other lineage in its range and which
evolves separately from all lineages outside its range.”
Let us take a look at Mayr’s definition, “populations that are reproductively isolated from other such groups” and let us try to understand
what it means and the effect of isolation.
REPRODUCTIVE ISOLATING MECHANISMS

A. Pre-zygotic isolation mechanisms prevent fertilization and zygote formation.

1. Geographic or ecological or habitat isolation – potential mates occupy different areas or habitats thus, they never come in contact. If two
populations of flies exist in the same geographical area, but one group lives in the soil and another lives on the surface of the water, members
of the two populations are very unlikely to meet and reproduce.

2. Temporal or seasonal isolation – different groups may not be reproductively mature at the same season, or month or year. Time is the
barrier that prevents species from interbreeding and producing sterile hybrids. Timing of the day when they are sexually active, the best
example will be that of the two fruit fly species. Drosophila persimilis and Drosophila pseudoobscura. The D. persimilis species are generally
active in the early morning, D. pseduobscura is active in the afternoon.

3. Behavioral isolation – patterns of courtship are different. For example, male fireflies of a variety of species signal to their female
counterparts by flashing their lights in specific patterns. Females will only respond to the signals flashed by their own species, preventing
them from mating with other closely related firefly species.

4. Mechanical isolation – differences in reproductive organs prevent successful interbreeding. It is caused by structures or that keep species
isolated from one another. For example, in flowering plants, the shape of the flower will tend to match up with a natural pollinator. Plants that
do not have the correct shape for the pollinator will not receive a pollen transfer.

5. Gametic isolation – incompatibilities between egg and sperm prevent fertilization. There is a couple of possible reasons why the egg and
sperm cannot unite in cases of gametic isolation. First, sperm and eggs have specific proteins on their surfaces that allow the sperm to
recognize the egg (and vice versa) and these proteins differ from species to species. So, if two different species mate, the sperm may be
unable to recognize the egg. Another example of gametic isolation happens when the sperm is unable to survive or will be less mobile in the
reproductive tract of a female from a different species.

B. Post-zygotic isolation mechanisms allow fertilization but nonviable or weak or sterile hybrids are formed.

1. Hybrid inviability – fertilized egg fails to develop past the early embryonic stages. When the zygote does form, but does not do so
completely and usually dies early in its life cycle.

2. Hybrid sterility – hybrids are sterile because gonads develop abnormally or there is abnormal segregation of chromosomes during meiosis.
When the zygote does grow to become a hybrid or the offspring of two different species of organisms, but the hybrid itself is unable to breed.

3. Hybrid breakdown - F1 hybrids are normal, vigorous and viable, but F2 contains many weak or sterile individuals. The hybrid is able to
breed, but future generations are not viable, sterile, or both.

Separate groups of organisms belonging to the same species may adapt in different ways to better exploit diverse environments or resources.
They also may evolve varied characteristics for attracting mates. That is, different groups evolve in different directions. Over time, these
groups or populations may become so different that they can no longer breed together--separate species are formed. One species does not
"turn into" another or several other species -- not in an instant, anyway. The evolutionary process of speciation is how one population of a
species changes over time to the point where that population is distinct and can no longer interbreed with the "parent" population. In order for
one population to diverge enough from another to become a new species, there needs to be something to keep the populations from mixing.
Often, a physical boundary divides the species into two (or more) populations and keeps them from interbreeding. If separated for long
enough and presented with sufficiently varied environmental conditions, each population takes its own distinct evolutionary path. Evolution
does not stop once a species becomes a species. Every population of living organisms is undergoing some sort of evolution, though the
degree and speed of the process varies greatly from one group to another. Populations that experience a major change in environmental
conditions, whether that change comes in the form of a new predator or a new island to disperse to, evolve much more quickly than do
populations in a more stable set of conditions. This is because evolution is driven by natural selection, and because when the environment
changes, selective pressures change, favoring one portion of the population more heavily than it was favored before the change.

Speciation is how a new kind of plant or animal species is created. Speciation occurs when a group within a species separates from other
members of its species and develops its own unique characteristics.

MODES OF SPECIATION

1. Allopatric speciation or geographic speciation (allo – other, patric – place; ‘other place’) - occurs when some members of a population
become geographically separated from the other members thereby preventing gene flow. Examples of geographic barriers are bodies of water
and mountain ranges.

2. Peripatric speciation. As in allopatric speciation, physical barriers make it impossible for members of the groups to interbreed with one
another. The main difference between allopatric speciation and peripatric speciation is that in peripatric speciation, one group is much smaller
than the other. Unique characteristics of the smaller groups are passed on to future generations of the group, making those traits more
common among that group and distinguishing it from the others.

3. Parapatric speciation (para – beside, patric – place; ‘beside each other’) – occurs when the groups that evolved to be separate species are
geographic neighbors. Gene flow occurs but with great distances is reduced. There is also abrupt change in the environment over a
geographic border and strong disruptive selection must also happen.

4. Sympatric speciation (sym – same, patric – place; ‘same place’) - occurs when members of a population that initially occupy the same
habitat within the same range diverge into two or more different species. It involves abrupt genetic changes that quickly lead to the
reproductive isolation of a group of individuals. Example is change in chromosome number (polyploidization).

No. 7
Devolopment of Evolutionary Thought
Douglas Futuyma defines evolution as “a change in the properties of groups of organisms over the course of generations…it embraces
everything from slight changes in the proportions of different forms of a gene within a population to the alterations that led from the earliest
organism to dinosaurs, bees, oaks, and humans.” “Evolution may be defined as any net directional change or any cumulative change in the
characteristics of organisms or populations over many generations—in other words, descent with modification… It explicitly includes the
origin as well as the spread of alleles, variants, trait values, or character states” by John Endler and in a very different vein, Leigh van Valen
characterized evolution as “the control of development by ecology;” this anticipates those who emphasize the importance of development in
evolution, including proponents of “evo-devo”.

EVOLUTIONARY THOUGHTS CONTRIBUTORS

Carl Linnaeus was the famous 18th century Swedish botanist and naturalist who created the basic biological taxonomy — the so-called
binomial classification system — that is the foundation of our modern taxonomic system. Linnaeus’ classification, at its most basic, uses the
dual "genus, species," nomenclature to classify organisms — everything from slime molds and bacteria to elephants and humans. When
Linnaeus finished his doctorate, he published a brief pamphlet that would eventually revolutionize the fields of biology and scientific
taxonomy. This "list" was written in Latin and was called Systema Naturae ("The System of Nature"). It proposed a radical new approach to
the ordering and classification of plants and animals. His system was hierarchically ranked, meaning that organisms were grouped into
successively larger groups based on morphological traits (that is, physical attributes). At the broadest level, the classification system was
divided into three broad kingdoms: animals, plants and minerals (the mineral designation was subsequently dropped). These categories were
further subdivided into increasingly specific designations, which included "classes," "orders," "genera," and "species." Linnaeus combined
two terms, genus and species, and used this combination to identify each particular organism. The species designation, a term he borrowed
from the English naturalist and parson John Ray, indicates the most basic unit of classification, traditionally defined as organisms capable of
interbreeding. The genus designation (gens is Latin for "tribe") ranks above species and designates the larger group of related organisms. For
example, a coyote (Canis latrans) is a different species from a wolf (Canis lupus), but both belong to the same genus, Canis. This genus, in
turn, could then be related to the higher-order ranks, such as order (Carnivora), class (Mammalia) and so on, all the way up to the highest
rank, the kingdom ranking (Animalia).

Thomas Robert Malthus was born near Guildford, Surrey in February 1766. Malthus' most well-known work, 'An Essay on the Principle of
Population' was published in 1798, although he was the author of many pamphlets and other longer tracts including 'An Inquiry into the
Nature and Progress of Rent' (1815) and 'Principles of Political Economy' (1820). Malthus first pointed out that human nature being what it
is, the passion between the sexes appears to be fairly constant and, if unchecked population will double itself every twenty-five years.
"Population, when unchecked, increases at a geometrical ratio. Subsistence increases only in an arithmetical ratio. A slight acquaintance with
numbers will show the immensity of the first power in comparison with the second." And this leads to Malthus’s principle of population.
Because of this unequal power between production and reproduction, "population must always be kept down to the level of the means of
subsistence.” While Malthus was not the first one to notice this, he was the first to inquire into the means by which this leveling of population
is achieved.

Georges Cuvier possessed one of the finest minds in history. Almost single-handedly, he founded vertebrate paleontology as a scientific
discipline and created the comparative method of organismal biology, an incredibly powerful tool. It was Cuvier who firmly established the
fact of the extinction of past life forms. He contributed an immense amount of research in vertebrate and invertebrate zoology and
paleontology, and also wrote and lectured on the history of science. Catastrophism was a theory developed by Cuvier based on
paleontological evidence in the Paris Basin. He observed something peculiar about the fossil record, instead of finding a continuous
succession of fossils, he noticed several gaps where all evidence of life would disappear and then abruptly reappear again after a notable
amount of time. Cuvier recognized these gaps in the fossil succession as mass extinction events. This led to the formation of Theory of
Catastrophism. It states that natural history has been punctuated by catastrophic events that altered that way life developed and rocks were
deposited.

James Hutton (1726–1797), a Scottish farmer and naturalist, is known as the founder of modern geology. He developed the theory of
gradualism according to which profound changes to the Earth, such as the Grand Canyon, are due to slow continues process and not part of
catastrophes as proposed by the Theory of Catastrophism. Hutton described a universe, that one formed by a continuous cycle in which rocks
and soil are washed into the sea, compacted into bedrock, forced up to the surface by volcanic processes, and eventually worn away into
sediment once again. Relying on the same methods as do modern field geologists, Hutton cited as evidence a cliff at nearby Siccar Point,
where the juxtaposition of vertical layers of gray shale and overlying horizontal layers of red sandstone could only be explained by the action
of stupendous forces over vast periods of time. There Hutton realized that the sediments now represented by the gray shale had, after
deposition, been uplifted, tilted, eroded away, and then covered by an ocean, from which the red sandstone was then deposited. The boundary
between the two rock types at Siccar Point is now called the Hutton Unconformity. The fundamental force, theorized Hutton, was
subterranean heat, as evidenced by the existence of hot springs and volcanoes.

Charles Lyell was a Scottish lawyer and the foremost geologist of his day. He is best known as the author of Principles of Geology. It has
popularized geologist James Hutton’s concept of “uniformitarianism” — the idea that the Earth was shaped by slow-moving forces still in
operation today. Uniformitarian ideas opposed the common belief among many geologists that unique catastrophes or supernatural events,
like the biblical flood in the story of Noah, shaped Earth’s surface. The motto of uniformitarianism was “the present is the key to the past.”
Lyell’s friend, Charles Darwin, took that idea and extended it to biology. Lyell argued that the formation of Earth's crust took place through
countless small changes occurring over vast periods of time, all according to known natural laws. His "uniformitarian" proposal was that the
forces molding the planet today have operated continuously throughout its history. He also wrongly assumed that these causes must have
acted only with the same intensities now observed, which would rule out asteroid impacts and the like.

Jean Baptiste Lamarck (1744-1829) is one of the best-known early evolutionists. Unlike Darwin, Lamarck believed that living things evolved
in a continuously upward direction, from dead matter, through simple to more complex forms, toward human "perfection."
Species didn't die out in extinctions, Lamarck claimed. Instead, they changed into other species. Since simple organisms exist alongside
complex "advanced" animals today, Lamarck thought they must be continually created by spontaneous generation. According to Lamarck,
organisms altered their behavior in response to environmental change. Their changed behavior, in turn, modified their organs, and their
offspring inherited those "improved" structures. For example, giraffes developed their elongated necks and front legs by generations of
browsing on high tree leaves. The exercise of stretching up to the leaves altered the neck and legs, and their offspring inherited these acquired
characteristics. Conversely, in Lamarck's view, a structure or organ would shrink or disappear if used less or not at all. Driven by these
heritable modifications, all organisms would become adapted to their environments as those environments changed. “The frequent use of any
organ, when confirmed by habit, increases the functions of that organ, leads to its development and endows it with a size and power which it
does not possess in animals which exercise it less.” Decreased use has the opposite effect: “The permanent disuse of an organ, arising from a
change of habits, causes a gradual shrinkage and ultimately the disappearance and even extinction of that organ” Another contribution of
Lamarck is the Theory of Inheritance of Acquired Characteristics. If an organism changes during life in order to adapt to its environment,
those changes are passed on to its offspring.

CHARLES DARWIN’S VOYAGE AND HIS OBSERVATIONS THAT LED HIM TO WRITE “THE ORIGIN OF SPECIES”

In 1831, Charles Darwin received an astounding invitation: to accompany Captain Robert FitzRoy of the HMS Beagle as ship's naturalist for
a trip around the world. For most of the next five years, the Beagle surveyed the coast of South America, leaving Darwin free to explore the
continent and islands, including the Galápagos. He filled dozens of notebooks with careful observations on animals, plants and geology, and
collected thousands of specimens, which he crated and sent home for further study. Darwin later called the Beagle voyage "by far the most
important event in my life," saying it "determined my whole career." When he set out, 22-year-old Darwin was a young university graduate,
still planning a career as a clergyman. By the time he returned, he was an established naturalist, well-known in London for the astonishing
collections he'd sent ahead. He had also grown from a promising observer into a probing theorist. The Beagle voyage would provide Darwin
with a lifetime of experiences to ponder—and the seeds of a theory he would work on for the rest of his life.

San Cristobal Island was the first island Charles Darwin visited arriving to the Galapagos on September 16th, 1835. The Beagle anchored in a
calm bay on the south of the island, near the actual capital of the Galapagos. The Beagle spent eight days surveying the coast. Darwin landed
five times pushed by his interest on the volcanic and cratered island. He studied carefully the lava flows and theorized about its formation.
His first impression of the dry coast he saw was of a deserted and isolated place. He did not find the tropical richness he expected, instead he
encountered desert plants, almost all-in flower, and some reptiles that he pointed on his notes. In this island he had the chance to collect the
first specie that later on would be the base and foundation of his Theory of Evolution, the San Cristobal Mockingbird. Floreana was the
second island explored by the Beagle expedition. Here Darwin had three days to collect species and the second bird to lead him to important
conclusions in the future, the Floreana Mockingbird. He realized the difference in between the previous specimen found in San Cristobal, and
started to pay more attention to this specie. Later he would discover the four species to be found in the archipelago and understand a pattern
of evolution due to the adaptation differences from each other according to the environmental features of each island. Isabela was the third
island to arrive in his voyage on September 29th, 1835. The trip around the island and through the channel in between Fernandina and Isabela
was noted on his field book. Darwin described the island as the most deserted and volcanically active. When sailing next to it, he observed
the lava flows and the smoke coming out from the craters. The Beagle decided to anchor in a place named Tagus Cove because of the easy
water bay this place forms. Darwin disembarked on October 1st and explored the volcanic terrain. Here he found the land of iguanas, both
marine and terrestrial that to him were ancient creatures that he describes more in Santiago visit. After Isabela, the boat sailed around Pinta,
Genovesa and Marchena, offering Darwin the chance to admire the different formations. Santiago was the last island were Darwin
disembarked on October 8th. This is the island where Darwin stayed the longest, by this time, he knew already that the islands were
something bigger and more important than they seemed when he first arrived. On his visit he expends 2 weeks and walked the whole island
with some crew members that helped him carrying the specimens he was collecting. Here he first noticed the difference in between the
tortoises from different islands with his own eyes; he was impressed by the amount of tortoises, which had different shapes and sizes. He
wrote a big deal of characteristics of their behavior and also had the chance to try their meat in soup. In this island Darwin noticed that most
species were similar but different from other in the other islands, giving enough evidence to theorize that species change and this is related to
their feeding and surroundings. He collected finches that helped him to understand this resolution. These animals are now considered the
world’s fastest evolving birds because of the adaptations they rapidly developed to cope with their needs in such a changing environment.

Charles Darwin’s theory argued that organisms gradually evolve through a process he called “natural selection.” In natural selection,
organisms with genetic variations that suit their environment tend to propagate more descendants than organisms of the same species that
lack the variation, thus influencing the overall genetic makeup of the species. Darwin, who was influenced by the work of French naturalist
Jean-Baptiste de Lamarck and the English economist Thomas Malthus, acquired most of the evidence for his theory during a five-year
surveying expedition aboard the HMS Beagle in the 1830s. Visiting such diverse places as the Galapagos Islands and New Zealand, Darwin
acquired an intimate knowledge of the flora, fauna, and geology of many lands. This information, along with his studies in variation and
interbreeding after returning to England, proved invaluable in the development of his theory of organic evolution. Darwin had formulated his
theory of natural selection by 1844, but he was wary to reveal his thesis to the public because it so obviously contradicted the biblical account
of creation. In 1858, with Darwin still remaining silent about his findings, the British naturalist Alfred Russel Wallace independently
published a paper that essentially summarized his theory. Darwin and Wallace gave a joint lecture on evolution before the Linnean Society of
London in July 1858, and Darwin prepared On the Origin of Species by Means of Natural Selection for publication. Published on November
24, 1859, Origin of Species sold out immediately. Most scientists quickly embraced the theory that solved so many puzzles of biological
science, but orthodox Christians condemned the work as heresy.