Lamarck: The theory of transformation

Though he was building on the work of his mentor, Count George-Louis Leclerc de Buffon,
Jean-Baptiste Lamarck (1744-1829) is often credited with making the first large advance toward
modern evolutionary theory because he was the first to propose a mechanism by which the
gradual change of species might take place. Also, he extended the definition of the change over
time, saying that life started out simple and became more complex. In 1809 he published
Philosophie Zoologique, in which he described a two part mechanism by which change was
gradually introduced into the species and passed down through generations. His theory is
alternatively referred to as the theory of transformation or simply Lamarckism. Though today
Lamarck's work is considered a major step forward, in his lifetime he did not receive much
recognition.

Use and Disuse

 Figure%: Use and disuse in the evolution of the neck of the giraffe

The classic example used to explain the concept of use and disuse is the elongated neck of the
giraffe. According to Lamarck's theory, a given giraffe could, over a lifetime of straining to reach
high branches, develop an elongated neck. A major downfall of his theory was that he could not
explain how this might happen, though he discussed a "natural tendency toward perfection."
Another example Lamarck used was the toes of water birds. He proposed that from years of
straining their toes to swim through water, these birds gained elongated, webbed toes to better
their swimming.

These two examples demonstrate how use could change a trait. By the same token, Lamarck
believed that disuse would cause a trait to become reduced. The wings of penguins, for example,
would be smaller than those of other birds because penguins do not use them to fly.

Lamarckian Inheritance

The second part of Lamarck's mechanism for evolution involved the inheritance of acquired
traits. He believed that traits changed or acquired over an individual's lifetime could be passed
down to its offspring. Giraffes that had acquired long necks would have offspring with long
necks rather than the short necks their parents were born with. This type of inheritance,
sometimes called Lamarckian inheritance, has since been disproved by the discovery of
hereditary genetics.

An extension of Lamarck's ideas of inheritance that has stood the test of time, however, is the
idea that evolutionary change takes place gradually and constantly. He studied ancient seashells
and noticed that the older they were, the simpler they appeared. From this, he concluded that
species started out simple and consistently moved toward complexity, or, as he termed it, closer
to perfection.

What are the stages of man evolution?

Any documentation identifying the stages of man evolution has to assume that man evolved rather than having
been created. We believe that evidence has failed to support the evolution from ape to man or any other type of
macroevolution.

The Cambrian explosion and the complete absence of transitional fossils testify against evolution. The fossil record
shows all life forms appearing fully formed and not changing during their tenure on earth, except for extinctions.
This information, and the recent finding that human DNA is losing its vitality by developing genetic disorders
supports devolution, the opposite of evolution.

The story below represents how evolutionists describe the stages of man evolution. The timeframe for the stages
of man evolution from the ancestor of both man and the modern ape to modern man is not known, but I will give
you an abbreviated chronology of what has been discovered from fossil remains over the years.

First of all, the word, hominidae, is used to describe the total member species of the human family that
have lived since the last common ancestor of both man and the apes. A hominid is an individual species
within that family, and the field of science that studies the human fossil record is known as
paleoanthropology. It is made up of two disciplines of paleontology, which is the study of ancient life
forms, and anthropology, which is the study of humans. Each hominid name consists of a genus name
(Australopithecus, Homo) which is always capitalized, and a species name (africanus, erectus) which is
always in lower case.

To begin our study of the stages of man evolution, the earliest fossil hominid, Ardipithecus ramidus, is a
recent discovery dating 4.4 million years ago. He was 4 feet tall and bipedal (having two feet). It is thought
this species lived as forest dwellers. Australopithecus anamensis, a new species, was named in 1995 and
was found in Kenya. This species lived between 4.2 and 3.9 million years ago, and its body showed
advanced bipedal features, but the skull closely resembled the ancient apes.

Australopithecus afarensis lived between 3.9 and 3.0 million years ago. He retained the apelike face with a
sloping forehead, a ridge over the eyes, flat nose, and a chinless lower jaw, and height, 36 and 5 tall. He
was fully bipedal, and the thickness of his bones showed he was quite strong. His build was similar to a
human, but the head and face were proportionately much larger.

The Australopithecus africanus was similar to the afarensis, but lived between three and two million years
ago. He was also bipedal and slightly larger in body size. His brain was not advanced for speech. The
hominid was an herbivore and ate tough, hard to chew, plants. The shape of the jaw was human-like.

The Australopithecus robustus lived between two and 1.5 million years ago. His body was similar to that of
the africanus, but had a larger and more massive skull and teeth. His huge face was flat and had no
forehead. He had no indication of speech capabilities.

The Australopithecus boisei lived between 2.1 and 1.1 million years ago. He was smaller than the robustus,
but with a more massive face. He had huge molars, for which the largest measured 0.9 inches across. Some
authorities believe the robustus and boisei are of the same species.

Next is the Homo habilis, or also called The Handy Man because tools were found with his fossil remains.
He existed between 2.4 and 1.5 million years ago. The brain shape shows evidence some speech had
developed. He was 5 tall and weighed about 100 pounds.

Homo erectus lived between 1.8 million and 300,000 years ago. Toward the end, his brain was that of the
size of modern man, and definitely could speak.

Erectus developed tools, weapons, fire, and learned to cook his own food. He traveled out of Africa into
China and the Southeast Asia developing clothing for northern climates. He turned to hunting for his food,
and only his head and face differed from modern man. Homo sapiens (archaic) lived during the period
200,000 to 500,000 years ago. He had speech capabilities; his skull was rounded with smaller features. The
skeleton shows a stronger build than modern human, but well proportioned.

Homo sapiens neandertalensis lived in Europe and the Mideast between 150,000 and 35,000 years ago.
 Brain size averaged larger than modern man, but the head was shaped differently, longer and lower. His
nose was large and extremely different from modern man in structure. He was a massive man, about 56
tall with a heavy skeleton that showed attachments for massive muscles. He was far stronger than modern
man, and his jaw was massive with a receding forehead like erectus.

Homo sapiens sapiens first appeared about 120,000 years ago, which is our own species.

Although this sounds like quite a convincing story, it is very slim on evidence and big on evolutionary
presupposition. What is not included in the above story is a long list of frauds, deceptions. and extrapolations from a
few small bone fragments into complete descriptions of semi-human or human life forms.

Dr. Joseph Mastropaolo draws the following conclusion. The only scientific objective, valid, reliable, calibrated
studies, which any biologist may verify, prove all so-called ape-men are frauds or forgeries. To date, no contrary
evidence has overturned these scientific objective, valid, reliable, calibrated studies. Therefore, the ape-man
alternative to Adam and Eve is based upon the anti-biology of frauds and forgeries whereas the historical and Eve
are based upon sound biology.

Genetics
Mendelian Genetics

When Gregor Mendel began his hybridization experiments with pea plants in 1856,
knowledge of how heredity works was limited. If two organisms of different height produced
offspring, it was assumed that the offspring's height would be somewhere between the
height of the two parents. This notion of blending inheritance presented a significant
obstacle for the acceptance of the theory of natural selection, since variation would be
removed from a population by being blended into nonexistence.

However, for some characteristics -- discrete traits -- inheritance did not produce a state of
being between the parents. The children of a brown-eyed father and blue-eyed mother do
not end up with an intermediate eye color; rather, children inherit the eye color of a single
parent. It was with these types of characteristics that Mendel performed his famous
botanical experiments. After carefully selecting pea plants to breed true for particular traits,
he then cross-bred strains with conflicting phenotypes (observable physical characteristics).
Most importantly for those who were to follow him, he meticulously catalogued the results of
these experiments.

Meiosis and Mitosis

The processes of meiosis (the production of sex cells) and mitosis (cell division) are both
integral operations necessary to an organism's survival and reproduction. Mitotic divisions
occur throughout the body; the results of mitotic division are two daughter cells, each
containing a diploid set of chromosomes. Meiosis, however, only occurs within the sex cells
of an organism. This process initially divide one cell into two, each with a diploid set of
chromosomes. However, this cell generation divides once again, without a corresponding
division of DNA, resulting in the final product - the egg or sperm - containing a haploid
(single) set of chromosomes. Additionally, during the process of meiosis, homologous
chromosomes may exchange material in a genetic recombination event called crossing over.
Crossing over is one of the ways that, with each subsequent generation, genes are
constantly reshuffled. The reformulation of novel combinations produces the variation
necessary for natural selection to operate.

Follow these links to view animated demonstrations of meiosis and mitosis.

Mendel's Experiments

From his experimental results, Mendel inferred that inheritance must occur via discrete
"particles", one from each parent, and that some of these characters are dominant -- that
is, preferentially expressed -- over others. The major insights attributed to Mendel are often
summarized in two "laws":

Principle of Segregation: observable traits are passed down to offspring via

(discrete, non-blending) particles, one from each parent.

Principle of Independent Assortment: the particles (later dubbed genes by

American geneticist T. H. Morgan) from each parent are equally likely to be passed
down to offspring.

The relationship between pea plant phenotype and genotype was carefully manipulated by
Mendel, who inferred the above principles through assiduous, empirical observation of trait
distribution.

How could he have figured out that there were two factors involved in inheritance, and that
one preferentially expressed itself over the other? If we assume that each parent
contributes a single chromosome to his/her offspring for a given trait, then we can calculate
the probable variants found in the first generation of offspring:

First Hybrid Generation

Yellow Parent's Traits

Y Y

Yg Yg
g
.25 .25
Green Parent's Traits
Yg Yg
g
.25 .25

All of the offspring from a cross (mating) between YY and gg parents should be alike in their
outward appearance (Yg). This is indeed what Mendel observed as all of his first generation
pea plants produced yellow seeds; however, it is a cross between the first-generation
offspring that provides some additional, critical information:

Second Hybrid Generation

 First Parent's Traits

Y g

YY Yg
Y
.25 .25
Second Parent's Traits
Yg gg
g
.25 .25

The final phenotypic proportions (3 yellow and 1 green) differ from the second-generation's
genotype frequencies (1 : 2 : 1 for YY : Yg : gg). This disparity between genotype and
phenotype provided the information necessary for Mendel to propose that some genes are
dominant while others are recessive.

# of Yellow Phenotype # of Green Phenotype

Pea Hybridization Generation
(Genotype) (Genotype)

1 Yellow 1 Green
F0 (parents)
(YY) (gg)

All Yellow 0 Green

F1
(Yg) (Yg)

3 Yellow 1 Green
F2
(1 YY, 2 Yg) (1 gg)

Besides stimulating an entirely new field of study, Mendel's work provides additional support
for Darwin's theories. The tenets of Mendelian genetics would prove to be instrumental in
supporting the concept of evolution by natural selection, allowing independent variation to
be preserved over many generations through hidden variation; additionally, this variation
could be recombined in innumerable novel combinations in future generations. Darwin's
problem explaining "blending" and the preservation of variation was essentially solved.
However, while Mendelian genetics provided a way to refute arguments concerning the lack
of a mechanism for the preservation of variation through inheritance, Mendel's results went
largely unnoticed until 1900, when the laws were independently "rediscovered" by several
geneticists -- Hugo de Vries, Carl Correns, and Erich von Tschermak -- some 34 years after
Mendel's original publication of his findings in 1866! (Click here to see a copy of Mendel's
original paper from 1866.) It would not be until the early 1930s that a modern
understanding of the relationship between genetics, morphology, and evolution became
integrated into what is now called the Modern Synthesis (Ch. 3).

If you are interested in trying some experiments with Mendelian hybridization but don't
have the time to catch fruit flies, give the Virtual Fly Lab a visit!
Molecular Genetics

Now that we know genes exist, how is it that genes impact living organisms? This was (and
still is) a fundamental question which has no complete answer. However, what is known is
that all of the processes which govern an organism's growth, development, and
maintenance are ultimately based in a code located in its chromosomes.

All cells that constitute an organism contain a nucleus, where the genetic material called
deoxyribonucleic acid, or DNA resides. The structure of the DNA molecule resembles a
twisted ladder and is often described as a "double-helix" (pictured at left). The vertical
sections are composed of sugar and phosphate molecules that provide structural support for
the molecule. The "rungs" of the double-helix ladder are composed of four different types of
molecules called bases. These bases -- adenine, thymine, guanine, and cytosine -- have
very specific bond affinities; adenine will only bond with thymine, and guanine exclusively
with cytosine. So, for example, a given sequence of connected DNA base-pairs might look
like the following:

...ATTACGGATAAT...

...TAATGCCTATTA...

and so forth. The information encoded in the base-pair sequence is used

for functions such as protein synthesis within the cell. The actual process of
protein synthesis includes two stages: transcription and translation. During
transcription, the DNA molecule is unzipped and "read" by a type of RNA
(ribonucleic acid) molecule called messenger RNA (mRNA). A
complementary copy is then made of one of the exposed DNA segments, except that RNA
substitutes the base uracil for the DNA base thymine. During translation, the mRNA
molecule is decoded in triplets by transfer RNA (tRNA), which carry one of 20 amino acids.
Many of the 64 possible combinations of base triplets redundantly code for a single amino
acid - for example, GCT, GCC, GCA, and GCG all code for the amino acid alanine.
Additionally, there are a few triplet sequences which begin and end this process of protein
synthesis but do not code for any amino acid.

Chapter 3: The Modern Synthesis

The early interpretation of Mendelian genetics focused upon the discrete changes which
were produced by the underlying genetic structures. During the early 1900's this
perspective weakened earlier acceptance of evolution by natural selection. If change
primarily acted through discrete mechanisms, there is no basis for the gradual accumulation
of change necessary for natural selection to operate. However, it was through the work of
three biologists - J. B. S. Haldane, Sewall Wright, and Ronald A. Fisher - that the integration
of a theory of inheritance, the maintenance of variation, and a basis for the genetics of
continuous variation were applied to the evolutionary paradigm. These ideas were
subsequently expanded and refined by later researchers such as geneticist Theodosius
Dobzhansky, biologist Ernst Mayr, and paleontologist George Gaylord Simpson.
Population Genetics

Evolution can be described in many ways; one way is look at evolutionary change as a
change in genotype frequencies over time. If organisms are considered evolutionarily
successful if they have more offspring, and offspring are created from genes, then changes
in gene frequencies (or more specifically, genotype frequencies) will reflect successful
evolutionary phenotypes. Researchers in the field of population genetics examine
populations in terms of differing proportions of particular genotypes in order to determine
what, if any, evolutionary forces are active in that population.

Hardy-Weinberg Equilibrium

One of the theoretical tools utilized by population geneticists today was independently
developed in 1908 by a British mathematician named G. H. Hardy, and a German physician
named W. Weinberg. The Hardy-Weinberg equation expresses an ideal distribution of
genotypes within a population, assuming that the gene frequencies are known. The validity
of the results from Hardy-Weinberg analysis is contingent upon five factors, all of which
must be in effect :

1. no mutation

2. infinitely large population

3. random mating

4. no migration

5. no genetic drift

If all parameters are met, the genotype frequencies should emerge in constant proportions
based on the individual gene frequencies. This is because the above parameters work to
keep the probability of a particular gene being passed on to the next generation at a very
statistically predictable level. Therefore, by calculating the probability of each possible
genotype based on the contribution of each parent, the frequency of each genotype can be
predicted. Supposing two alleles - A and a - are present in a population where frequency(A)
= p and frequency(a) = q, the possible genotypes should occur in the following proportions:

Genes from parent 1

A = .5 a = .5

Genes from parent 2 A = .5 f(AA) = .25 f(aA) = .25

a = .5 f(Aa) = .25 f(aa) = .25

 The proportions of the three possible genotypes - AA, Aa, and aa - are 1 : 2 : 1

The Hardy-Weinberg Equilibrium can therefore be expressed as the equation, p2 +

2pq + q2 = 1, where p = f(A) and q = f(a)

A glance at the above parameters suggests that it is very unlikely they will all be fulfilled
within any given natural population. However, as with other theoretical notions based on
ideal principles, the main product of these calculations is to provide a comparison by which
a population geneticist can assess why the sampled population is not conforming to ideal
behavior.

Example 1

In a population of lab-bred flies, a gene controlling eye color is discovered. The R allele
produces regular colored eye pigment, while the r allele produces red pigment. Individuals
that are heterozygous (Rr) have pink eyes. In a population of 150 flies, 15 flies have red
eyes, 90 have normal eye color, and 45 have pink eyes. Check if this population is in Hardy-
Weinberg equilibrium.

Step 1: Determine gene frequencies

Phenotype Genotype # of Individuals

Normal Eyes RR 90

Red Eyes rr 15

Pink Eyes Rr 45

Given this information, calculating the allele frequencies is simply a matter of counting up all
of the alleles.

Remember, each parent carries two alleles, so the total # of alleles twice the
population.

Also remember that heterozygous individuals carry one of each allele.

Taking these two factors into account,

f(R) = [(90*2)+(45] / 300 = 225/300 = 0.75

f(r) = [(15*2)+(45)] / 300 = 75/300 = 0.25

Step 2: Determine expected genotype frequencies

Plugging the frequencies of each allele into the Hardy-Weinberg equation, we find the
expected numbers of each genotype in the population:

f(RR) = p2 = f(R)*f(R) = 0.5625

f(rr) = q2 = f(r)*f(r) = 0.0625

f(Rr) = 2pq = 2*[f(R)*f(r)] = 0.375

Multiplying each of these genotype frequencies with the total population number, we find
that there should be:

84 normal-eyes flies (AA)

9 red-eyed flies (aa)

56 pink-eyed flies (Aa)

Since partial individuals do not exist, the numbers are rounded off.
Step 3: Compare with original population numbers

Comparing the expected numbers with the actual numbers of each phenotype, population
geneticists can determine if populations are either in equilibrium (or very close to it) or are
experiencing disequilibrium of some sort. In this example:

Phenotype Genotype Expected # Observed #

Normal Eyes RR 84 90

Red Eyes rr 9 15

Pink Eyes Rr 56 45

In this example, the population is not in equilibrium since the expected and observed values
do not match. Disequilibrium can be attributed to different possible mechanisms, depending
on (1) the context of the population, and (2) the manner in which the population is skewed.

Disequilibrium

Disequilibrium refers to a difference between observed and expected ratios of genotypes

within a given population as we saw above. This situation may come about as a result of
small population size (which accentuates the effects of genetic drift), migration, and
selective forces which favor or hinder particular phenotypes. Additionally, disequilibrium can
arise in situations of non-random mating. What needs to interpreted are the relative
proportions of each genotype (and their corresponding phenotypes), how these compare
with expected values, and the elements of the environment which could promulgate
evolutionary changes.
For instance, in the above example of fruit flies, the lower incidence of pink-eyed flies
suggests that there may be selective forces acting against their survival, or perhaps
encouraging both homozygous conditions over the heterozygous one. Or perhaps the flies
are more likely to select mates with the same eye color, leading to a slight reduction of
heterozygous individuals each generation.

Hidden Variation

One of the phenomena which the study of population helps us to understand is the
preservation of variation at the genetic level. One example of this is the artificial breeding
and selection of all of the modern species of dogs from a wolflike ancestor. All of the genetic
variation necessary to produce phenotypes like the Chihuahua and the St. Bernard can be
found in the wolf genome. This hidden variation is due to the effects of polygenic effects of
genes, where many different loci additively affect a particular trait (for example, height). A
similar line of reasoning can apply to the question of why recessive lethal alleles linger in
population gene pools. This is due to the fact that most recessive alleles (lethal or not) are
not expressed in every generation; instead, they are stored by heterozygous individuals.

This is also the reason why inbreeding carries with it an increased likelihood of the
expression of a deleterious or lethal allele. It is very likely that all individuals carry some
deleterious alleles which are left unexpressed from generation to generation, primarily
because the frequencies of these alleles in the whole population is very low. While
outbreeding (or exogamy) will significantly reduce the probability of a chance encounter
between two individuals carrying the same deleterious alleles, inbreeding will significantly
increase this probability on the basis of degree of relatedness.

Chapter 4: Speciation and Phylogeny

What is a species?

The term species is defined as a group of organisms which interbreeds under natural
circumstances, producing viable, fertile offspring, and which is reproductively isolated from
other groups. This definition is known as the Biological Species Concept. One way of
evaluating this concept is to consider that a species will experience gene flow, which will
tend to maintain genetic compatibility between members. Conversely, organisms that do not
exchange genetic information - either through geographic or behavioral isolation - will
experience genetic drift, and tend to become increasingly different over time.

Speciation

How are species formed? Geographic isolation probably is the most obvious explanation, but
other possible modes include:

Allopatric speciation: geographic isolation which impedes gene flow between two
groups in a population (e.g. a mountain range, river, etc.)
 Parapatric speciation: partial geographic isolation, coupled with selective
pressures, that maintains species boundaries even with some gene flow.

Sympatric speciation: high selection pressures that create species boundaries

without geographic boundaries.

The Ecological Species Concept emphasizes the role of selection in maintaining species
boundaries, rather than purely abiding by the strict rules of allopatricity (total geographic
isolation).

Phylogeny

Taxonomy, the classification of biological species, is a system used to organize all of the
forms of life found on the planet of which we are aware. Biological taxonomic classification is
based upon a hierarchical system created by Carolus Linnaeus in the 18th century. However,
taxonomies can be arranged according to different criteria. This may be particularly
problematic in the realm of biological classification, since arbitrary bases for classification
would impede communication between researchers using different systems. For this reason,
biological classification has been based on phylogeny, or relationship by common ancestry.
Because any given group of organisms can only share a single common ancestor, this
system provides an adequately discriminating lingua franca for biologists to use throughout
the world.

The question of how phylogenies can be accurately determined, however, presents a difficult
problem. This field - determining the method and criteria used in classifications - is referred
to as systematics. There are several different schools of thought that take different
approaches to addressing these challenges.

phenetics: This approach emphasizes overall morphological similarity between

organisms by using large numbers of traits to assess and comparing these statistical
levels with other groups. One branch of this is referred to as numerical taxonomy

cladistics: This approach emphasizes relationships based solely on common

descent.

evolutionary systematics: This school bases taxonomies on both overall similarity

and common descent.

While cladistics provides a sound theoretical basis for constructing taxonomies, the
determination of common descent becomes increasingly difficult when applied to the natural
world, due to homoplasy, or evolutionary convergence. Cladists have developed a
methodology specifically to address this issue.
Reconstructing Phylogenies

Character states (see below) are the core component of phylogenetic reconstructions; more
specifically, it is character states relative to other groups with known phylogenies that make
phylogeny construction possible. This method differs from phenetics in that it is not quantity
of characteristics which leads to validity of analysis. Rather, the most important issue is
resolving variable traits into three classes, of which only one type is phyogenetically
informative.

symplesiomorphy: shared ancestral trait that is not present in all descendant

groups. An example of this is egg-laying, which is present in chickens and
monotremes (duck-billed platypus), but absent in humans and primates. These
characters do not provide useful information for reconstructing phylogenies.

autapomorphy: uniquely derived trait not shared with any other groups. An
example of this is the enlarged third digit of the aye-aye, Daubentonia
madagascariensis, which is not possessed by any other living primates. These
characters do not provide useful information for determining phylogenies either, but
are good for identifying particular species.

synapomorphy: shared, derived character shared by all descendants, to the

exclusion of a common ancestor. An example of this is live birth and lactation, which
characterizes all placental mammals, to the exclusion of other groups. These are the
only characters which can be used to reconstruct phylogenies.

As stated above, problems arise from the convergence of characters,

which may be incorrectly designated as synapomorphies. Familiar
examples of this is the identification of aquatic and aerial mammals
on the basis of homologous features of their physiology. Pictured at
the left are the dugong (top), the mole (middle), and the bat
(bottom). The dugong has forelimbs designed to assist paddling in
the sea, the mole has forelimbs specialized to dig in the ground, and
the bat has forelimbs designed for aerial maneuvers. However,
common descent is based on recognizing that each bone in the
forelimb is derived from a common ancestral structure for all three
animals.

So how do researchers decide which characters should be classified as synapomorphies?

This question hinges upon the issue of character polarity - which trait state is ancestral
versus which is derived. To accomplish this, a taxonomic group that is equally distant to all
of the groups in question must be incorporated into the analysis. This group is called the
outgroup. In the picture shown at the right, the group consisting of siamangs and gibbons
(Hylobatidae) could be used as an outgroup for comparing character states within the great
apes. The character state of the gibbons would be assumed to be more representative of
the ancestral condition. However, this technique is not entirely sound, for there is no clear
reason why we should assume the gibbons would have stopped evolving since their
divergence with the great apes. For this reason, character polarity is also augmented by
information gleaned from studies of ontogeny (embryology and development) and fossil
material, when available. Even with this repertoire of tools by which to create phylogenetic
trees, controversies abound; systematics remains one of the most contentious fields in the
biological sciences.

An excellent explanation of cladistic methods and terminology is at the University of

California Museum of Paleontology. Also be sure to check out the UCMP glossary of
phylogenetic terms for some quick definitions. After you've become more familiar with how
to classify based on phylogeny, try your hand at a systematics exercise for PC or Macintosh
computers from the University of Illinois.

Introduction to the Primates

Meet Your Relatives

Why study primates? By studying primates, we're essentially studying ourselves. The
primate order includes humans and our closest living biological relatives,as well as all
extinct primate and human ancestors. In addition to providing evolutionary insights into the
physiological and behavioral evolution of the human lineage, primates exhibit an
extraordinarily diverse array of behaviors and social systems, which allows them to exploit
many habitats within the tropics, ranging from savanna-woodland to rain forest. Some
species, such as the Japanese macaque (Macaca fuscata), have adapted to the snowy
winters of the island of Hokkaido. The rather cosmopolitan status that the primate lineage
boasts is one of a number of reasons to study their adaptive strategies in the context of
evolution. Finally, the cognitive capacity of primates provides a window into the evolution of
intelligence, providing deeper insights into the machinery that drives our own behavior and
thought processes.

Common characteristics of primates

The following characters distinguish the primate order from other mammalian groups. While
all primates do not necessarily have every one of these traits, all primates have trait
complexes that include a preponderance of the traits listed below. This is because some
groups have lost some of these traits or developed others over the course of their
evolutionary history.

1. opposable thumb and big toe

 o assists in grasping and manipulation behaviors

o adaptation to arboreal lifestyle

2. flat nails instead of claws, with dermatoglyphs (fingerprints) on fingers and toes

3. hindlimb-dominated locomotion

4. relative reduction in the olfactory sensory system (smaller snouts) as compared to

other mammalian orders

5. increased reliance on visual sensation

o eyes are large and exhibit a high degree of frontation, or placement toward
the front of the face

o frontation increases overlap of visual fields, increasing binocular vision

o each sends visual information to both hemispheres of the brain, enhancing

depth perception and producing stereoscopic vision

6. tendency toward smaller litter size, longer gestation times, and extended period of
juvenile growth

o increased period of maternal investment and care

7. relatively large brains

8. reduced number of teeth, with a maximum of two incisors, one canine, three
premolars, and three molars in each jaw quadrant

Taxonomy of Living Primates

The primate order is generally subdivided into four groups, which are colloquially the
prosimians, New World monkeys, Old World monkeys, and apes. However, different
divisions of the Primate order are produced by different systematic approaches--namely, the
differential weighting of overall similarity versus phylogeny. Two taxonomic arrangements
are generally used today--one based on the traditional division between Prosimii and
Anthropoidea, and a more recent one that divides groups into the Strepsirhini and
Haplorhini. The difference between these two taxonomies is that the Tarsiiformes have been
moved from the traditional suborder of Prosimii into the Anthropoidea. For the purposes of
this site, the traditional taxonomy consisting of a Prosimii-Anthropoidea division will be
used.

Suborder Infraorder Superfamily Family Subfamily Prosimii

(prosimians) Lemuriformes Lemuroidea
(lemurs) Cheirogaleidae
(dwarf and mouse lemurs) Lemuridae Lemurinae
(true lemurs) Lepilemurinae
(sportive lemurs) Indriidae
(indris) Daubentoniidae
(aye-aye) Lorisiformes Lorisoidea Lorisidae Lorisinae Tarsiiformes Tarsioidea Tarsiidae
(tarsiers) Anthropoidea
(anthropoids) Platyrrhini Ceboidea
(New World monkeys) Cebidae Cebinae
(e.g., capuchins, squirrel monkeys) Aotinae
(e.g., owl monkeys) Atelinae
(e.g., spider monkeys) Alouattinae
(e.g., howler monkeys) Pithecinae
(e.g., saki, uakari) Callimiconinae
(e.g., callimico) Callitrichidae
(e.g., tamarins, marmosets) Catarrhini Cercopithecoidea
(Old World monkeys) Cercopithecidae Cercopithecinae
(e.g., macaques, guenons, vervets) Colobinae
(e.g., colobus, langurs) Hominoidea
(apes and humans) Hylobatidae Hylobatinae
(e.g., gibbons and siamangs) Pongidae Ponginae
(great apes; e.g., gorilla, chimpanzee, and orangutan) Hominidae Homininae
(humans) Source: R. Martin, 1992: Classification of Primates, in S. Jones, R. Martin, and D.
Pilbeam, 1994, The Cambridge Encyclopedia of Human Evolution, Cambridge University
Press, Cambridge.

The Prosimians

The Prosimians are divided into three groups: Lemuriformes, Lorisiformes, and Tarsiiformes.
Lemurs (Lemuriformes) can only be found on Madagascar and the Comoro Islands, and
represent a lineage of primates that have evolved in isolation from other such groups over
the past 120 million years. Lemurs are primarily nocturnal, although some species (most
notably Lemur catta) are active during the day, or during portions of both day and night.

The continental prosimians consist largely of Lorisiformes, which can be divided into two
groups:galagos and lorises (pictured above, middle). The lorises are all nocturnal, and exist
in African and Asian forest regions. Their diets consist primarily of fruits, gums/exudates,
and insects.
Tarsiers are the third group of Prosimians. They are unique in their entirely faunivorous
dietary composition; additionally, they are particularly specialized for a vertical clinging and
leaping style of locomotion. Tarsiers can be found in Southeast Asia and Indonesia.

The Anthropoids

The Anthropoids are divided into two large groups along geographic lines: Platyrrhini (New
World primates) and Catarrhini (Old World primates).

Platyrrhines
There are two families within the New World monkeys:

The callitrichids (Callitrichidae) consist of marmosets and tamarins, which are small-bodied
primates that live in Central and South American rain forests. Several unique characteristics
include their dentition (they have one fewer molar in each quadrant than other
Anthropoids), the fact that they have clawlike nails instead of flat nails, and habitual
"twinning," i.e., females frequently give birth to twins. In addition to these physiological
characteristics, callitrichids also live in polyandrous groupings (i.e., single female with
more than one male).

The second family, Cebidae, consists of six subfamilies that encompass a wide range of
ecological and social diversity. This group also includes the only noctural Anthropoid, the owl
monkey or night monkey; additionally, several species of cebids also possess prehensile
tails. The cebids are all primarily arboreal, and exhibit a number of different social systems.

Catarrhines
The catarrhine primates are generally larger than platyrrhines; additionally, aspects of the
physiology (e.g., catarrhines have only two premolars in each quadrant vs. three for
platyrrhines) and ecology (e.g., catarrhines are found in many terrestrial habitats) differ
between these groups. Catarrhines can be divided into two superfamilies, the
Cercopithecoidea (Old World monkeys) and the Hominoidea (apes).

The cercopithecoids have two general groups, which can be divided along ecological and
dietary lines: the Colobine monkeys are distinguished by their specialized stomachs, which
have been modified for a highly folivorous, or leaf-eating, diet. Additionally, Colobines tend
to live in single-male, multifemale social groupings. Cercopithecines, the other goup within
the Cercopithecoidea, are composed of a cohort of monkeys that exhibit a wide range of
dietary, ecological, and social preferences. There are many terrestrial species, and their
geographic distribution includes Africa as well as Asia (and in one case, Gibraltar as well).

The apes represent a tailless group of large-bodied primates that proliferated during the
warm, moist Miocene epoch. The classification Hominoidea includes both lesser (e.g.,
gibbon, siamang) and great apes (orangutan, gorilla, chimp, human). The apes are
generally allocated into one of three families: Hylobatidae includes the lesser apes;
Pongidae includes the great apes, minus humans; Hominidae includes humans and their
fossil relatives.

The Earliest Hominids

A Very Brief History

The etymology of the word Australopithecus is, "southern ape," which harkens back to the
name given to the original specimen. In 1924, Raymond Dart examined the strange,
immature skull of a small-brained creature extracted from the cave breccia of a South
African mine. This was the first discovery of an Australopithecine, and was named Taung,
after the site where it was found; its species designation Dart gave it was apt--
Australopithecus africanus, or "southern ape from Africa". The acknowledgement of Taung
as a member of the human lineage remained hotly contested for many years after the initial
finding, primarily because of preconceived notions of which characteristics embodied the
human precursor. Also, many researchers of Dart's time were convinced that the human line
started in Asia and not Africa, despite the paucity of evidence from both regions.
Additionally, with the previous discoveries of a fossil "hominid" called Eoanthopus dawsonii,
the phylogenetic position of Taung (as well as many other fossils) remained questionable. It
was not until 1953 that E. dawsonii, more commonly known as the Piltdown Man, was
exposed as a scientific fraud (to read more about the Piltdown hoax, click here).

The Piltdown hoax did much to reveal preconceptions and underlying motives within the
study of human origins. It too perfectly resolved a primary issue - placing the evolution of
the brain ahead of locomotory developments - and remains one of the most interesting case
studies of the scientific establishment. However, after the discovery of the hoax, once again
our origins had to be refigured.

Introducing the Australopithecines

Ardipithecus ramidus

The earliest fossil hominid - while not classified as an Australopithecine - reaches back in
time to 4.4 Mya. This specimen, discovered in 1992 by Tim White (University of California at
Berkeley) off the Middle Awash River in Ethiopia, has been variably reported as an ancestor
to all hominids, or the ancestor to hominids and modern chimpanzees.

Australopithecus anamensis

The genus Australopithecus spans a portion of geologic time from approximately 4.2 Mya to
about 1.0 Mya. The earliest form, a species called Australopithecus anamensis, was only
recently announced by Meave Leakey from the Kenya National Museum (1994). Dated from
between 4.2 and 3.5 Mya, these fossils exhibit characteristics of the postcranium which
suggest that A. anamensis was bipedal, although dental and mandibular morphology
indicates a more apelike physical disposition. These fossils have been found in the western
Lake Turkana region in northern Kenya.

A. afarensis

The most famous specimen of Australopithecus - "Lucy" - belongs to the species, A.

afarensis. Found at sites in East Africa (e.g., Laetoli, Hadar), A. afarensis - which spans a
time range between 3.8-2.8 Mya - has provided a great deal of fossil information concerning
the morphology of early hominids. Most of the characteristics are continuous, and their
condition within A. afarensis lies somewhere between other hominids (relatively lower), and
chimpanzees (relatively higher):

subnasal prognathism (degree of facial projection beyond the borders of the

braincase whe viewed in profile)

canine size dimorphism

diastema

endocranial capacity (approximately 404 cc, which is very close to modern

chimpanzees)

compound temporal-nuchal crests

sagittal crest

V-shaped dental arcade

These traits are variably expressed across later Australopithecine species.

Later Australopithecines

The remaining groups of Australopithecines can be divided along two lines: geographic
distribution, and general morphological features. By geography, these hominids can be
subdivided into South African and East African groups; along morphological lines, they can
be grouped as either gracile (lightly built), or robust (heavily built).

Spatiotemporal Distribution of Australopithecines

Species Geographic Location Temporal Distribution Morphology

A. africanus South Africa 3.6 - 2.2 Mya gracile

A. robustus South Africa 2.0 - 1.4 Mya robust

A. boisei East Africa 2.2 - 1.2 Mya robust

 A. aethiopicus East Africa 2.5 - 2.2 Mya robust

South African Forms

A. africanus

A. africanus includes both Sts 5 (named "Mrs. Ples"), and the previously mentioned Taung
child. A. africanus is very similar to A. afarensis in postcranial anatomy, with elements of the
pelvis, ribs, and vertebrae exhibiting similar characteristics. The cranium of A. africanus is
somewhat different, with a higher degree of cranial flexion and no crests (sagittal or
compound T-N), as observed in A. afarensis. These specimens have been found
predominantly at the South African sites of Sterkfontein and Makapansgat, in addition to
Taung.

A. robustus

A. robustus is also within the genus Australopithecus; however, these fossils differ markedly
from the more gracile forms. Their crania exhibit remarkably well-fortified structures and
buttresses to counteract increased forces in mastication (chewing). Some of these
characteristics include a large sagittal crest, as well as flaring zygomatic arches
(cheekbones), both of which provide additional leverage and anchor points for the muscles
of mastication. Additionally, the teeth of A. robustus are very large, particularly in
comparison to the anterior (front) dentition.

East African Forms

A. aethiopicus

A. aethiopicus, discovered by Alan Walker (Penn State University), stirred quite a

controversy over the phylogenetic arrangement of the hominid lineage. Until this discovery,
most phylogenies had the Australopithecines evolving down one exclusive lineage, while
Homo evolved separately down another (although both shared a common ancestor,
presumably A. afarensis). A major portion of this puzzle included a tight relatioship between
the robust Australopithecines, an arrangement which required rethinking because of the A.
aethiopicus specimen, KNM-ER 17000 (aptly named the Black Skull). Because it bore a
number of similarities with A. afarensis which suggest its placement as an intermediate form
between A. afarensis and A. boisei, the previous arrangement with the South African A.
robustus became questionable. Were the South African A. robustus fossils still part of a
common lineage including A. boisei, or could South and East African Australopithecines have
evolved independently of each other?

A. boisei

A. boisei, the East African robust form, is thought to have descended from A. aethiopicus.
This species exhibits a hyper-robust morphology, with exaggerated features similar in kind
to those of A. aethiopicus, but far greater in degree. The massive dentition is coupled with a
sagittal crest; increased dishing of the face, or orthognathism, characterizes these forms.
This reduction in postnasal facial projection probably came as a result of increased demands
to combat masticatory forces. Specimens include OH 5 and KNM-ER 406, among others.
Whereas earlier interpretations of their dental specialization offered dietary shifts (e.g.,
consuming a tougher, lower quality diet), recent analyses of dental microwear suggest that
A. boisei incorporated a diverse range of foods in its diet.

Early Homo

The first specimens of a different type of hominid emerged approximately 2.3 Mya in East
Africa. These hominids exhibited a different suite of characteristics, including:

higher cranial capacity

reduction in tooth size (particularly molars and premolars)

reduction in masticatory apparatus

parabolic dental arcade

thinner tooth enamel than Australopithecine species

less prognathic facial profile

These specimens have been traditionally attributed to Homo habilis,

although recent investivators have alleged that perhaps two (or more)
species are represented in the assemblage commonly called H. habilis.

In addition to the above characteristics, early Homo also appears

concurrently with a stone tool technology known as the Oldowon
Industry.

Bipedal Locomotion

The principle feature which binds all hominids together into a unique group is their choice of
locomotion. All hominids exhibit features of their anatomy which indicate habitual
bipedalism. There are a great many features which provide direct or subtle evidence for
this behavior; a some of the main features of the skeletal system include:

shape and proportioning of the pelvic girdle

positioning of the head and neck of the femur

position of the big toe (abducted, as opposed to divergent)

position of the foramen magnum, the hole on the underside of the cranium which
transmits the brainstem and spinal cord
Other features include asymmetry of the distal femoral condyles and proximal tibial
condyles; relative positioning of the ischium and the iliac crest; angle of the femoral neck.
Additionally, reconfiguration of muscular attachments, especially the hip abductors, is
required to maintain proper balance while moving in a bipedal gait. These characteristics,
when observed in fossilized skeletal elements, argue for an organism which utilized a highly
specialized mode of locomotion shared with all human beings today.

Theories on the Origin of Bipedality

Two researchers from UC Davis - Peter Rodman and Henry

McHenry - examined the locomotor efficiency of bipedalism. They
found bouth bipedalism and quadrupedalism to be similar in energy
Locomotor
costs; however, examining out closest living relatives (the Great
efficiency
Apes) suggests that perhaps early hominids adopted a similar
suspensory posture, from which the move to bipedalism could be
facilitated.

Peter Wheeler (Liverpool John Moores university) suggested that an

erect posture was adopted by early hominids to combat high
temperatures, especially when travelling exposed to direct sunlight
Thermoregulation (as in open savanna). The benefits of bipedalism would be twofold:
reduction of exposed area of the body to sunlight, along with an
increased exposure to cooling winds (Click here to see a drawing of
this).

The benefits of being able to transport things manually are

obvious. Some species of primates have adapted mechanisms to
Freeing hands for perform these tasks (e.g., cheek pouches in Cercopithecines);
other uses however, the ability to carry a resource either to save for later, or
to consume in a safer location, would provide allow for flexibility in
planning for early hominids.

Kevin Hunt (Indiana University) recently proposed that the

emergence of bipedalism came as a postural feeding adaptation.
Observing common chimpanzees in the wild, he determined that,
although they rarely walk bipedally, when they choose to adopt a
Postural feeding bipedal position it is almost always in the context of feeding on fruit
from small trees. This increases foraging efficiency, particularly
when fruiting trees are spread about; feeding chimpanzees can
shuffle over to the next tree, rather than get down on all fours, got
to the next tree, and get back up.

A new member of the family - A. garhi

Found at and around a site called Bouri, in Ethiopia, this species was only recently named.
The general morphology of this hominid is similar to the other Australopithecines, though A.
garhi has a sagittal crest and their canines are generally larger. And surprisingly, there is
evidence that this species of Australopithecine used stone tools, a trait generally associated
only with the genus Homo. The discovery of this species prompts researchers to ask new
questions about the lineage leading to Homo sapiens sapiens, since it is dated at just about
the time to be the ancestor of the first Homo species.

For more information on this new Australopithecine, click here, and to keep up on more
recent developments in paleoanthropology, click here.

Click here for a comparison of numerous hominid (and great ape) crania. Or visit an online
resource for fossil hominids - a very handy list with nicknames, museum specimen numbers,
descriptions, and links.