2A Evolution

02/08/2019 Evolution
ORIGIN OF LIFE
Origin of Life - 3.8 billion years ago, Earth was formed 4.5 billion years ago.
Life can be defined as an entity that can carry out self sustaining activities on its own, respond to environment and reproduce, and is capable of Darwinian
evolution
Origin of life only occurred once, diversity of life is due to evolution.
Features of living entities
Chemical complexity: Cellular-Molecular organisation, Biochemical Pathways, Genes and Gene expression, Molecular Genetic Pathways
Microscopic Organisation
Self Replication, Self Assembly
Mechanism for sensing and responding to the environment
Capable of evolution
Biochemical Abiogenesis
Generation of life from non-living matter

Life has progressed through a route of increasing biological complexity, starting from simple inorganic molecules
Early atmosphere had Hydrogen, Ammonia, Methane, Carbon Dioxide, Water and Nitrogen. But no Oxygen
These gases combined in different proportions to produce bio-organic molecules
Oparin One celled life forms came from simple organic molecules
Life arose on earth when early atmosphere was subjected to energy inputs. (Heat, UV)
Haldane
Lightning could have provided sudden but large inputs of energy for attaining complexity
Spark passed in a chamber containing a mixture of gases resembling primordial atmosphere - Carbon Dioxide, Ammonia,
Water, Nitrogen, Hydrogen, No Oxygen
Led to evidence of purines, pyrimidines, some other organic molecules
Abiotic synthesis of ribose and of nucleosides is much more difficult
Miller Supramolecular polymerisation of molecules could not be attained
The experiment has had very little success in replication over the years
Nitrites are formed which makes the water acidic, and prevents formation of amino acids
According to Jeffrey Bada, primitive earth also had Iron and Carbonate minerals which neutralised nitrites and acids, and
allowed formation of amino acids
Energy released from redox reactions led to synthesis of organic molecules and formation of oligomers and polymers.
Gunter Wachterhouser
But silent on the availability of catalysts.
Clay Origin: Complex organic molecules arose on a non-organic replication platform.

Graham Cairns Smith
Silicate crystals in combination with Ni-Fe provided a catalytic system.
Prebiotic soup led to formation of short RNA molecules.

RNA can store genetic information and function as a catalyst. RNA could be used as a template for synthesis of peptides in
an auto-catalytic reaction
Peptides came to play an increased role in RNA replication
Co-evolution of RNA and proteins
Led to evolution of a primitive translation system with RNA genome and RNA Peptide catalysts.
Led to Cooperation and Competition among RNA
Led to evolution of complex nucleic acid structures, and ultimately the DNA genome.
RNA world
Prebionts: Non living supra molecular structures that later evolved into the first living cells.
Coacervates: Organic molecules surrounded by a film of water molecules. They selectively absorb materials from
surrounding water and incorporate them into their structure. Represent well-organised, self replicating systems
Protobionts: Organic molecules surrounded by a double membrane. Can capture, transform and utilise energy.
Functional and structural specialisation in earliest cells
Emergence of Eukaryotic cells by Endosymbiosis
Early divergence of Eukaryotes
Emergence of Multicellular forms
Growing diversity due to speciation.
THEORIES OF EVOLUTION
Lamarck Inheritance of Acquired Characters
Changes in environment make special demands on organisms — leading to use and disuse of organs, or need of novel
organs
Use and disuse of organs affects their efficiency, leading to new characteristic traits. e.g. Appendix in humans are vestigial,
but they are used in whales; Evolution of a long neck in Giraffe.
Characteristics acquired by the parents during their lifetime can be transmitted through reproduction to offspring.
Variations over several generations accumulate to form new species.
Lamarck suggested that no species went extinct. Over time, they just became different by acquiring novel traits.
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Limitations:
Ernst Haeckel showed that not all traits acquired during lifetime are transmitted to the next generation. e.g. Cutting off tails
of rats.
Neo Lamarckism - Morgan and Cope: Acquired characters which become incorporated in the germ plasm are heritable. These
acquired characters accumulate over generations and lead to formation of new species.
Variation: Individuals in a population have inherent variation

Limited Resources: Therefore, struggle for existence leads to only a fraction of offspring surviving in each generation. If
all individuals that are born reproduce successfully, population size would increase exponentially.
Survival is determined by characters that fit the changing environment the best. Fitter individuals leave behind more
offspring. The least fit individuals do not contribute offspring to the next generation.
Over generations, it leads to a gradual change in population composition, with favourable characteristics accumulating over
the generations.
Gradual changes lead to formation of new species
Darwin
Natural Selection
Essentially, NS is differential success in reproduction.

NS operates through an interaction between the environment and inherent variability in the population.
Helpful variations are 'selected for' i.e. they are passed on to the next generations.
It leads to adaptation of organisms to their local environment, and over generations contributes to formation of distinct
species, due to accumulation of variations.
Survival of the fittest (Herbert Spencer): Organisms with favourable variations will survive, because they are the fittest to face
their surroundings.
New species originate as a result of large, discontinuous variations which appear suddenly due to mutations
Mutants are markedly distinct from their parents
Mutations are distinct and larger in effect than the continuous variations, as propounded by Darwin.
Mutations may occur in any direction, and unsuitable mutations get weeded out by NS
The incipient stages have no importance in speciation overall.
Hugo De Vries: Mutation
Theory
Criticism
Results were based on studies in Primrose. Cytological studies showed that the mutants were mostly polyploid mutants,
instead of gene mutants. Polyploidy is a common occurrence in plants.
Morgan: Mutations in Drosophila were of all magnitudes. Mutations alone don’t account for evolution. Rather, they furnish
the raw material for evolution.
Synthesis Theory
Combines several findings from cytology, morphology, genetics and classical Darwinian and Lamarckian theories, with
Population Genetics. (Darwin + Mendel)
Pioneers of the theory: Dobzhansky (genetics), Ernst Mayr (species), George Simpson, Wright and Fischer (statistical
basis)
Evolution is viewed as a change in allele frequencies within populations.
Components
Biotic Potential: Innate capacity of population to increase under optimal conditions, with suitable age and sex ratios
present.
Variations
Hereditary Variations: important for evolution as they can be passed on to offspring.
Non hereditary Variations: appears during life of an animal and are not passed on to offspring
Evolution begins at the population level. The factors that lead to changes in allele frequency are Selection, Mutation,
Drift and Gene Flow.
Genetic Drift is the chance fluctuation in allele frequencies. It is significant in smaller populations.
NS selects for fitter variants in each generation, causing an increase in frequency of genotypes in proportion to their fitness.
Genetically controlled variations are produced by Mutations, Chromosomal Change, Recombination. These are the raw
material on which NS acts. While mutation creates novel genetic material, recombination creates different combinations
from this novel material.
Gene flow due to migration also affects allele frequencies.
Micro Evolution: Smaller changes below the species level. Largely indiscernible at the phenotypic level and mostly does
not involve reproductive isolation.
Macro Evolution: Changes at the level of species or genera, accompanied by reproductive isolation.
HARDY WEINBERG EQUILIBRIUM
A population in HW equilibrium is NOT evolving

Represents a quantitative approach in evolutionary analysis
Population’s genotype and allele frequencies will remain unchanged over successive generations
Enables us to compare a population’s actual genetic structure over time. If genotype frequencies vary from that expected under HW equilibrium, it means one
of the model’s assumptions are being violated.
P + Q = 1 — Allele Frequencies

P^2 + 2PQ + Q^2 = 1 — Genotype Frequencies
Deviations from HW
Assumptions
Non Random Mating
Population is large
Sexual Selection
No gene flow between populations
Breeding Territories, Courtship Displays, Chemical signaling
Mutations are negligible
Consanguineous mating
Individuals are mating randomly
Assortative mating — preference to phenotypically similar which leads to homozygote excess
NS is negligible
Dis-assortative mating — preference to phenotypically dis-similar which leads to heterozygote excess
NATURAL SELECTION
Natural Selection is the process by which favourable heritable traits become more common in successive generations of a population of reproducing organisms,
as a result of differential fitness of genotypes.
NS acts on a phenotype and improves its survival, thereby selecting for the underlying genotype.
NS occurs through an interaction between the environment and the variability inherent in a population
NS — Adaptation — Niche Specialisation — Speciation.
According to Darwin, NS is the agent that leads to survival of the fittest
Selection can act at any level of biological organisation and at any stage of life.
Viability Selection: Survival until adulthood

Sexual Selection: Success determines who will parent the next
generation
Survival Selection: Longer survival in reproductive phase increases
the number of offspring.
Fecundity Selection: Limits fecundity
Gametic Selection: Difference in viability of gametes
Compatibility Selection: Union of some combination of egg and
sperm might be more compatible.
Stabilising Selection Directional Selection Disruptive Selection
Favours a single phenotype and allele

Favours extreme traits over intermediate trait
frequency shifts towards extreme values of
Selection works to maintain genetic values.
that phenotype.
polymorphisms. Variance increases as population diverges into
Advantageous allele will increase in
It maintains stable frequencies of multiple two extremes.
frequency independent of its dominance
alleles Happens in case of adaptive radiations arising
relative to the other allele. Even if it is
When a certain value of a trait has much from a single population, with a given trait
recessive, it is eventually fixed.
higher fitness than all other values diverging into multiple fitness maxima
e.g. selection towards darker morphs
corresponding to different niches.
in Biston betularia moths
Continuous Variations Discontinuous Variations
Variations are very small and in a graded series. Genetically quantitative

Sudden, large
and change between generations is small.
Rare
Polygenically controlled.
New and conspicuous characters appear suddenly and spontaneously
Subjected to directional, stabilising and disruptive selection
NS acts rather strongly to select for or against
NS acts weaker than for discontinuous variation.
According to Darwin, plays a minor role in speciation.
According to Darwin, plays a significant role in speciation
MUTATIONS
Stable and heritable changes in genetic material

Sudden, random and heritable. May be beneficial, harmful or neutral
Enrich the existing gene pool of a population
Mutations lead to intra-specific phenotypic differences. As members within a species are not exactly alike, NS affects them differentially.
Hugo De Vries first used the term mutation to describe sudden heritable phenotypic changes in evening primrose.
Muller first used X Rays to induce mutations in a fruit fly.
Most of the mutations are inconsequential because
Large volume of genome is transcriptionally inactive DNA. Hence, a mutation in these regions does not have any effect on phenotype
Silent or Synonymous mutation: Due to codon degeneracy, a mutation might not lead to coding of a different amino acid. Hence, inspite of a change in
genotype, there is no alteration in phenotype.
Mutation in various theories
Lamarck: No mention of mutation. Environmental forces leading to new characters however is analogous to the concept of mutations
Darwin: Continuous variations are more significant in speciation than discontinuous variation.
Bateson: Contradicted Darwin, discontinuous variations play a bigger role in speciation.
De Vries: Mutation as the driving force behind origin of new species.
At the level of an individual base
Base substitution: Transition or Transversion

Point Mutations
Frameshift: Insertion or Deletion
Point Mutation: Silent (codon for same AA), Mis sense (codon for another AA), Non sense (changed to stop codon)
At the level of chromosome
Structural
Amplifications
Increased dosage of genes
Due to slipped strand phenomenon and unequal cross over events
Multiple copies of chromosomal regions
Triplet Expansion: Triplet codon is amplified in several copies
Gross Mutations Deletion: Mostly leads to harmful variations
Bring separate genes in proximity: Translocations, Interstitial deletions, Inversions
Loss of Heterozygosity
Numerical
Aneuploidy: Only few chromosomes are affected. Leads to diseases like Turner’s Syndrome, Down syndrome and Klinefelter
Syndrome
Polyploidy: Involves an entire chromosomal set. Mostly due to meiotic failure. Often leads to formation of new species in plants.
Mechanism of Mutation
Spontaneous Mutation: Tautomerism, Depurination, Deamination
Induced Mutation
Radiation: Ionising or non Ionising
Heat
Chemical Mutagens: Base analogs, alkylating agent, intercalating agent
Role of Mutations in Speciation
Anagenesis: Accumulation of changes gradually transforms a species. Mutations are of fundamental importance in anagenesis
Cladogenesis: Splitting of gene pool into two or more separate pools. When gene pools separate, mutations accumulate differently in each population,
reinforcing the reproductive isolation.
Mutations lead to adaptive radiations, which reinforces speciation
Polyploidy: Mutations that leads to instant speciation.
GENETIC DRIFT
Random, non-directional process of change in allele frequencies

One of the basic mechanism of evolution along with mutation and selection.
Acts independent of selection. Leads to non-adaptive evolution
Significant in small populations, not as important in larger populations where other agents have a stronger effect
May cause loss of genetic diversity, because it may cause certain variants to disappear (more likely for the rarer alleles)
Two sets of small populations isolated from one another may differ significantly due to drift
Genetic Drift Natural Selection
Random and non-directional process Directional process

Doesn’t occur due to environment challenges Due to environmental challenges
Non adaptive Adaptive
Important alleles may disappear and harmful ones may get fixed Towards increasing frequency of fittest alleles
Operate on neutral alleles as well Does not operate on neutral allele
Bottleneck Effect Founder Effect
Reduction in population size. e.g. when a natural disaster kills a large proportion of the population, leaving
survivors in which allele frequency composition is very different from the original population
Consequent generations have very little genetic variation Small subset of population founds a
This new subset does not have simila
composition as the original populatio
SPECIATION
Species is a group of populations whose members can interbreed with one another under natural conditions and produce fertile offspring, and are
reproductively isolated from other such species.
Speciation occurs when members of similar populations no longer interbreed to produce fertile offspring within their natural environment. Groups in species
diverge and become reproductively isolated.
A single species can lead to formation of many different species.
Reproductive Isolation
Mechanisms that prevent two individuals in a species from mating and producing viable and fertile offspring. It is the prevention of gene flow.
Geographical: Populations cannot mate due to geographical barriers.

Pre Zygotic: Isolation before fertilisation
Ecological isolation — live in different habitats, and cannot come together
Temporal isolation — Breed at different times of the year, or day (day/night)
Behavioural isolation — Patterns of courtship are different. e.g. different songs in birds
Mechanical incompatibility — Reproductive organs are incompatible. e.g. in right handed and left handed shelled snails
Gametic isolation — Sperm of one species does not survive the internal environment of the female. Mating happens, but fertilisation fails.
Post Zygotic: Isolation after fertilisation
Hybrid Inviability
Hybrid infertility
Hybrid Breakdown: F1 hybrids may be viable and fertile, but consequent generations are not.
Allopatric Speciation that occurs in geographically isolated populations
Allopatric populations diverge because they accumulate different set of mutations, are subjected to differing selection pressures and
independently undergo genetic drift
Small sample of a large population becomes isolated at the periphery of the range
Peripatric
Similar to founder effect
Individuals are more likely to mate with those in their geographical vicinity.
There are no obvious barriers to reproduction. Mating behaviour is not random
Parapatric
There might be a hybrid zone where the two populations come in contact. But if Hybrid fitness is lesser, there is selection against the
hybrid, reinforcing isolation between the two populations.
Sympatric Species splits into two groups that diversify and become genetically isolated while remaining in the same place.
EVOLUTIONARY PATTERNS
Sequential Evolution
No new populations are formed

Descendant population is not genetically identical to its predecessor
Random fluctuations over a long period of time without producing new species.
Changes due to mutations, variations, natural selection and drift produce only temporary changes, which fluctuate at random
Changes are not directional
Divergent Evolution
Origin of new populations from old ones

Most evident form of evolution
What we see in fossil records is divergent evolution
Based on timescales (and correspondingly, divergence)
Microevolution: Subspecies/geographic races. Evolution in its simplest form operating at population level
Macroevolution: Evolution of species/genera
Megaevolution: Large scale evolution of families, orders, classes and phyla
Micro Evolution
Drift + Mutation + Selection + Gene Flow = Relatively Small changes in population

Change in gene frequencies
Changes genetic equilibrium in a Mendelian population
Operates below the species
Recombination also changes gene frequency in gene pool
Mutations can be accelerated by certain environmental and chemical factors
Offspring of a population are different genotypical and phenotypically due to action of NS
Changes produced in genotype may manifest at the phenotype level as positive or negative
Phenotype interacts with the environment, as a result of which unsuitable phenotypes and underlying genotypes diminish in frequency
Successional Microevolution: Within a single population, which results in replacement of pre-existing populations by the new ones. Can be seen in successive
strata of paleontological series.
Clines: Microevolution in response to gradual changes in climate.
Divergent Microevolution: Splitting of parental population into two or more new populations. Isolation operates as an additional factor. Due to isolation,
microevolution occurs independently in the separated populations.
Macro Evolution
Production of new adaptive types through population fragmentation and genetic divergence
Each sub population exhibits changes in a definitive adaptive direction
Macro evolution tends to occur in a group of individuals which have entered a new adaptive zone free of competition.
In a new adaptive zone, number of individuals is less and hence, resource competition is less intense.
Within the new adaptive zone again, each sub population can accumulate mutations and evolve independently (Parallel specialised adaptations)
e.g. Mammals entered a new niche and diverged independently in aquatic, terrestrial, arboreal habitats
Specialisation may lead to narrow adaptive sub zones. Over specialisation may lead to extinction, due to lack of ability to respond successfully to
perturbations.
Mega Evolution
Origins of new levels of biological organisation

E.g. Fishes — Amphibians — Reptiles
Parental stock develops some pre-adaptations, which enables them to enter a new zone, crossing ecological barriers
This entry into a new zone must be rapid, as extreme negative selection operates at the barrier level
Then, this population radiates in all directions and ceases to be a panmictic group anymore
Over time, evolution of specialised adaptations leads to reproductive incompatibility
Development of exoskeleton helped amphibians colonise land and led to first reptiles
From reptiles, both mammals and birds diverged
Common features of Macro and Mega Evolution
Pre adaptations (Not necessary in micro evolution)

Invasion of new zones (Micro evolution does not necessarily require a change in habitat)
Specialisation + Loss of evolutionary flexibility (in micro evolution, gene frequencies fluctuate and hence, flexibility remains)
Always divergent (micro evolution is not)
Phyletic Relationships
Monophyly: Taxa whose members have descended from a common ancestor

Polyphyly: Taxa whose members are descended from multiple ancestors
Paraphyly: Consists of groups MCRA and all descendants, except a few.
Character States
Apomorphy: Specialised character unique to a group or species (present in all species in the
clade, hence a diagnostic character)
Plesiomorphy: Character present in a few species in the clade but not in all (hence not a
diagnostic character)
Synapomorphy: Character shared by an ancestor and its descendants
Homoplasy: Character showed by a set of species but not in their MRCA
Stasigenesis: Lineages that neither split nor change but persist in the same manner for millions
of years. e.g. Sphenodon, Coelacanths
Based on rate of Evolution
Bradytelic Evolution: Gradualism. Slow evolution in minor ways

Tachytelic Evolution: Origin of species by abrupt changes. Intermediate fossil forms would then be absent (which is not the case)
Punctuated Equilibrium (Stephen Jay Gould): Most evolutionary changes are rapid bursts of speciation alternating with long periods of stasis
MOLECULAR DRIVE
Term coined by Gabriel Dover in 1982 to describe evolutionary processes that change the genetic composition of a population through DNA turnover
mechanisms (MOT)
Molecular drive operates independently of natural selection and genetic drift
Difference from Drift: Changes in the frequencies of the individual alleles that occur through its action are not random in their direction. If a certain population
of genetically identical organisms is divided into several smaller populations, then genetic drift will lead to fixation of different alleles in each population. In
contrast, the effect of molecular drive should lead to fixation of the same alleles in all populations.
Difference from selection: Alleles that are fixed through its action need not favourably affect the phenotype of the organism and can thus have a zero or even
negative impact on the biological fitness of the individual
In molecular drive, one allele is replaced by another not because this is more advantageous for its bearer, but because, at the level of the DNA, it multiplies
more effectively through mechanisms like gene conversion, unequal crossing-over, transposition, slippage replication and RNA-mediated exchanges
Entails a number of mechanisms connected primarily with replication, recombination and repairing of nucleic acids. These mechanisms favour the formation
and proliferation of certain sequential motifs in the gene pool of the population regardless of the degree to which the existence of these motifs is manifested in
the phenotype of the organism and the degree to which it affects its biological fitness
All MOT are essentially non-mendelian, in that the initial mutant sequence can increase or decrease in copy number within the lifetime of an individual
The best-known such process is the concerted evolution of genes present in many tandem copies, such as those for ribosomal RNAs
Proposed to extend to the diversification of multigene families
Because mutations changing the sequence of one copy are less common than deletions, duplications and replacement of one copy by another, the copies
gradually come to resemble each other much more than they would if they had been evolving independently.
Concerted evolution can be unbiased, in which case every version has an equal probability of being the one that replaces the others.
However, if the molecular events have any bias favouring one version of the sequence over others, that version will dominate the process and eventually
replace the others.
MIMICRY
Mimicry is physical or behavioural resemblance of a species to another species (which are not necessarily closely related taxonomically) or object in surrounding in
order to ward off or escape from threats such as predators.
Types of Mimicry
Protective
Concealing Mimicry e.g. Camouflage
Warning Mimicry
Batesian
Mullerian
Aggressive e.g. Ant mimicking spider
Conscious e.g. Death Feigning
Mimicry provides protection to the mimic
Concealing Mimicry: organisms search for a background that matches their colour, or changes their coloration to blend in the
surrounding. e.g. Stick Insect
Protective
Warning Mimicry: Harmless species resembles a harmful species, in order to ward off predators or enemies. e.g. King Snake
mimics coloration of Coral Snake.
Auto Mimicry: Occurs within a single species, where an animal mimics parts of its own body. e.g. False eyes at the back in
caterpillars, false head in butterflies.
Predator mimics a harmless model in order to gain advantage in capturing prey

Aggressive
e.g. Ant mimicking spider
Animal behaves as if they are dead.

Conscious e.g. American Opposum, Hognose Snake
Molecular surface markers of the virus resemble the host body’s protein so that the immune system treats the virus as self.
Evolution of Mimicry
Sudden mutation leads to phenotypic changes.

The great advantage conferred by this phenotypic change is strongly selected for, increasing frequency of mimics in the population. Over time, the non-mimics
get wiped out.
Batesian Mimicry Mullerian Mimicry
Protective mimicry in which species that is edible or harmless resembles

an inedible or harmful species, and is therefore avoided by predators. Two or more distasteful or poisonous organisms resemble each other.
e.g. Viceroy butterfly resembles the Monarch butterfly e.g. Bee and Wasp
Exploitation of model by mimic Mimicry Ring: Group of organisms all mimicking the same pattern.
Theoretically, selection favours the mimic only if it is less common than Mutualistic arrangement
the model. (Negatively Frequency Dependent)
FOSSILISATION
Fossil is naturally preserved evidence of a living organism directly (bones, body parts) or indirectly (impressions, foot marks).
Fossils are found mainly in sedimentary rocks.
Body Fossil: Actual parts like bones, shells, leaf imprints; Trace Fossils: Tracks, burrows, casts
Factors favourable for preservation
Presence of hard material in form of exoskeleton or endoskeleton. Soft tissues get decomposed or destroyed
Detachment from atmosphere after death as soon as possible. Otherwise, action of oxygen and microorganisms lead to decomposition. Temporary covering by
sand, silt or clay facilitates fossil formation. Because marine organisms are not exposed to atmosphere, marine fossils are usually better preserved.
Rapid sedimentation over the dead remains leads to compression and gradual hardening.
Environment: In acidic medium, calcareous skeletons are lost by dissolution. Siliceous skeletons are lost in alkaline medium.
Inorganic minerals in solution in ground water often replace organic skeletal material. This makes the fossil harder and preserves it better.
Less deformation and metamorphism due to orogenic movements, igneous intrusion etc. which may destroy well preserved fossils.
What do fossils tell us?
Movement: Tracks of insects, trails of worms, foot prints of vertebrates

Habitat: Burrows of worms, Tunnels of invertebrates
Reproduction: Eggs, Larval stages
Nutrition: Mouth arrangement, Faecal pellets
Significance of Fossils
Biostratigraphic classification of sedimentary rocks

Correlation and age determination
Indicators of past environment — Paleoecology, which may help us understand drivers for current scenario of climate change
Reconstruction of Paleogeography — continental drift
Evidence of prehistoric life, helps in understanding evolution
Understand past sedimentary environment.
Source of pre historic DNA
Limitations of Fossils
No hard parts are generally preserved. Soft parts are almost always damaged to some degree
Terrestrial organisms are mostly decomposed
Bottom of sea fossil — eaten by deposit feeders or scavengers, difficult to access
Do not provide comprehensive information. Difficult to piece together information from isolated fossil discoveries.
Dating Fossils
Carbon Dating
Once an animal or plant dies, it doesn’t take in C-14 anymore. C-14 present begins to decay.
In living beings, C-14 is constantly getting converted to N-14, so the mixture remains the same as the atmosphere.
By comparing C-14/C-12 ratio of present atmosphere and comparing it to that in the fossil, the age of fossil can be ascertained, by knowing the rate of decay of
C-14. (1/2 of C-14 will convert to N-14 in 5730 years)
Anything above 50,000 years old has very little 14C left in it.
Used only for organic material, cannot be used for rocks.
SYSTEMATICS
Species Concept
Morphological Species Concept (Aristotle, Linnaeus): Species are recognised by their essential characters expressed in terms of morphology
Cuvier (1829): Assemblage descended from one another or from common parents. Such a species concept disregards variation within populations. It proposes
the existence of a limited number of types.
Species consist of similar individuals
Each species is separated from all others by sharp discontinuity
Limitations: Variations in individuals belonging to same species (for e.g. sexual dimorphism), similar looking species are reproductively isolated.
Nominalistic Species Concept: Only individuals exist, while species are man’s own creation.
Biological Species Concept (Mayr): Species are groups of interbreeding natural populations that are reproductively isolated from other such groups.
Members of a species form a reproductive community
Species is a distinct ecological unit that interacts with other species
Species is a genetic unit that holds a large gene pool
Species is the smallest independently evolving unit. Each species has its evolutionary trajectory.
Limitations: Asexual reproduction, Parthenogenesis, Hybridisation
Evolutionary Species Concept: Species is a spatio-temporal lineage of populations that evolved separately from other lineages and has its own ecological
niche.
Phylogenetic Species Concept: Diagnosable cluster of individuals within which there is a parental pattern of ancestry and descent.
Exceptions to Species Concept
Evolutionary Intermediates: Organisms that may appear to be alike, but are different species because they do not interbreed.
Ring Species: Species with geographical distribution that forms a ring and overlaps at the ends. The different populations can interbreed with their adjacent
populations, but not with others.
Cryptic Species: Similar looking species, that do not interbreed
Chronospecies: Different stages in the same evolving lineage that existed at different time points.
Species classification is contentious, because species are dynamic, evolving classes but we attempt to force them into rigid classes.
But species classification is important, because it is the primary unit for study of biogeography, evolution, selection, adaptation, speciation, etc.
Principles of Classification
Classification is a system of naming objects or entities by common characteristics, such as structural/functional similarities or evolutionary relatedness.
(Taxonomy is the science of classification)
It is a process of establishing, defining and ranking taxa within hierarchical series of groups.
Assumption is that, greater the degree of similarity, closer the biological relationship
Taxonomy is the science of grouping biodiversity into species, describing the species, and classifying this diversity into higher level taxa, that reflect evolutionary
history.
Process of Classification
Begin by looking for anatomical features that appear to have the same function as those found on other species. Thus, classification is based on homology
(similarities that indicate shared ancestry).
Principle of Homonymy: Use of same name for different taxa must not occur.
Principle of Priority: If the several scientific names are given to a single animal by different scientists, the senior-most name is selected by law of priority. Rest
of the names are called junior synonyms
Taxa should be described in a phylogenetic context and should be monophyletic
Taxa should be strongly supported by measures such as bootstrapping, Bayesian Posterior Probability and congruence among independent datasets (like
molecular and morphological data)
Linnaean hierarchy of ranks should be used i.e. Kingdom, Phylum, Class, Order, Family, Genus, Species
Synonyms: Different names assigned to the same taxon should be mentioned along with the valid taxon
Names in Latin must be descriptive of the species.
Description must be made on the basis of a type specimen
Current scheme should be disturbed as little as possible
Empty or redundant categories must be minimised
Anatomical features of different organisms that have similar structure even if they are used for different functions, because they
were inherited from a common ancestor.
Homology More homologies two organisms possess, the more likely it is that they have a close genetic relationship.
e.g. arm of human and wing of bat
Homology could be biochemical as well e.g. DNA or Proteins
Anatomical features that have similar form and function, but no underlying evolutionary relatedness to a common ancestor
e.g. wings of bird and butterfly
Analogy
Convergent Evolution
Can confuse traditional classification based on homology
Issues in classification of organisms
Defining species boundaries, as speciation is a continuous process.

Sometimes closely related organisms may differ significantly on important characteristics
Frequent changes in name and assigning many synonyms affects stability.
Why are names of taxa changed?
Insights from molecular taxonomy, with an aim to ensure generic names are monophyletic
Previously a single species based on morphology, found to consist of multiple genetic lineages
Morphologically different groups assigned species status, found to be of the same genetic lineage
Resolve species complexes
Importance of Classification
Makes study of a wide variety of organisms easy

Uniform system throughout the world, in order to avoid confusion — helps to organise efforts (Unique, Universal, Stable)
Understand interrelationship among different groups of organisms
Basis for study of biogeography, selection, evolution
Applied biology such as wildlife management, agriculture, pest management , conservation biology
Type Specimens
Holotype: Single specimen on which description of the species is based

Allotype: Opposite sex to the holotype
Paratype: All remaining specimens after designation of the holotype
Syntype: If no holotype is designated and the author studies multiple specimens for the description
Lectotype: Syntype designated as holotype. Others the Paralectotypes
Neotype: If all type specimen are destroyed, a specimen that fits the description can be designated as the neotype
Binomial Nomenclature
Binomial nomenclature is the system of scientific names applied to taxonomic units
Each species is assigned a two part scientific name

Scientific name is always written in italics (or underlined)
First word is a generic name, which starts with a capital letter
Second word is a specific name, written in lower case. It may give information about the organism: what it looks like, where it is found or who
discovered it.
Developed by Carlos Linnaeus
Universally accepted, to ensure uniformity. Common names can often be misleading
Each organism has an unique name.
Levels of organisation are Kingdom, Phylum, Class, Order, Family, Genus, Species
International Code for Zoological Nomenclature
Main entry for the binomial is followed by the full surname of the author who published the classification. Comma is used between the author’s name and the
year of publication. There should not be any punctuation mark between the binomial and the name of the author
e.g. Paramaecium aurelia Ehrenberg, 1833
New Combination: If the species was assigned to a different genus than what it is assigned today
Amoeba proteus (Pallas, 1766) Leidy, 1875 — This means Pallas originally described the species in some other genus but Leidy transferred it to Amoeba
When used with a common name, binomial name is given in parentheses
Abbreviation 'sp.' Is used when actual specific name cannot or need not be specified.
Tautonymy is permissible in zoological nomenclature
Family name should be based on the type genus
Trinomial nomenclature adds the name of a subspecies (Panthera leo persica)
When species names are after a person, endings are in i, ii and ae. When it is a geographical place, the species ends in ensis or iensis
If the several scientific names are given to a single animal by different scientists, the senior-most name is selected by law of priority
CLADISTICS
Phylogenetic systematics is concerned with determining which features encapsulate evolutionary relatedness
Ancestral Feature: Similarity derived from common ancestor of the group

Derived Feature: Similarity that arose within the group
Clade: Group of organisms related by common descent. A clade is thus, a monophyletic group.
Greater number of synapomorphies between two species, the more recent the common ancestor and the more closely they are related.
Outgroup: Used to assign character polarity, Character states shown by the outgroup are assumed to be ancestral, other states are considered derived. Polarity
assignments are most effective when several different outgroups are used.
Branching Points: Point in the tree where novel characteristics first appear
Advantages
Time trees using calibrations

Study origin of species and episodes of change.
Objective taxonomy
Reconstruction of ancestral states
APPROACHES TO TAXONOMY
Anatomical features are encoded by the underlying genotype. Closely related species have similar anatomical features
(homologous)
Ontogeny captures phylogeny
Anatomical features are investigated using a light microscope. Ultrastructure can be studied using electron microscope
Anatomy Anatomy is mostly used to supplement observations from morphology
Comparative Anatomy establishes correspondence between body parts of different organisms — both correspondence of origin and
of function
Anatomy contributes to embryology, phylogenetic analysis and palaeontology
e.g. Skull analysis in classifying reptiles, dentition in mammals, vertebrae in fish
Chromosome number and structure — active field of study in primates

Position of centromere
Cytology
Meiotic behaviour sometimes shows heterozygosity of inversions
Especially important in plants where chromosome number is a major driver of speciation
Physiology reflects gene expression

Functional aspects of biological diversity
'Extreme’ examples can be studied to gain insights into evolutionary adaptation. e.g. water conservation in desert organisms,
oxygen use efficiency in high altitudes, pressure tolerance in deep waters
More suitable to differentiate between populations adapted to different environments
Physiology
Limitations
Can only be measured in living specimens — no fossils, no museum samples

Physiology is more likely to be adaptive than DNA — parallel and convergent evolution
Physiology is plastic and may give different parameters in different environments
Molecular Taxonomy
Phylogeny can be inferred from molecular data — DNA, RNA, Protein

The more recently two species branches from a common ancestor, the more similar their DNA
Rate of change in DNA over evolutionary timescales vary among different parts of the genome
Molecular data is more likely to reflect gene level changes, and be less affected by convergence and parallelism (which is common in morphological datasets)
Molecular Taxonomy has revised many traditional notions
In traditional taxonomy, crocodiles, snakes, lizards and other reptiles are grouped together into Reptilia
Molecular studies show that Crocodiles are more closely related to birds than to lizards or snakes
Thus, Class Reptilia is paraphyletic instead of monophyletic
Monophyletic Group: taxon that includes MRCA and all of its descendants.
Paraphyletic Group: Taxon that includes MRCA but not all its descendants
Polyphyletic Group: Species derived from more than one MRCA
Comparison of sequences in homologous regions. Prior research goes into identifying suitable homologous regions for different
kinds of research.
DNA
Both nuDNA and mtDNA can be used
DNA sequences are aligned and used to construct a phylogenetic tree
Protein sequences can be aligned and analysed like DNA

Proteins are likely to be less variable than DNA, as it does not encapsulate variations that result in degenerate codons
Proteins
Less effective in capturing background mutations that are synonymous, but effective in capturing mutations that lead to selection
on phenotype
Sequence of 6-8 pairs of DNA that binds to a given RE

DNA Restriction Different lengths of fragments are produced upon enzyme treatment (RFLP)
Can be used to determine heterozygosity, or to compare RE profiles between 2 species to check allele similarity
Allozymes are different molecular forms of an enzyme

Correspond to different alleles of a common gene
Allozymes
Detected using electrophoresis
Mostly used for population level analyses but can also be applied to closely related species
Regions of DNA with short tandem repeats

Microsatellites The polymorphism corresponds to number of repeats at the micro satellite loci
Mostly used for population level studies
Random Amplified Polymorphic DNA

RAPD
Mostly within species genetic studies, but can also be applied for closely related species
Amplified Fragment Length Polymorphism

AFLP
Electrophoresis separates the amplified DNA fragments that exhibit length polymorphism
Organisms are identified based on mtDNA.

Barcoding
Cytochrome Oxidase genes are most suitable — conserved within a species, but variable across species
Numerical Taxonomy
Numerical Taxonomy involves analysis of various types of taxonomic data by mathematical/computerized methods and numerical evaluation of the similarities or
affinities between taxonomic units, which are then arranged into taxa on the basis of their affinities
Construction of Taxonomic Groups

Individuals are selected and their characters spotted out. Larger the number of characters, better is the approach for generalization of the taxa
Resemblances among the individuals are then established on the basis of character analysis
Discrimination of the Taxonomic Groups
When the taxonomic groups chosen for the study show overlapping of characters, discrimination should be used to select them
Cluster Analysis is used to construct dendrograms
Seven principles of numerical taxonomy have been enumerated by Sneath and Sokal
1. Greater the content of information in the taxa, and more the characters are taken into consideration, the better a given classification system will be
2. Every character should be given equal weightage in creating new taxa
3. The overall similarity between any two entities is a function of the individual similarities in each of the many characters, which are considered for comparison
4. Correlation of characters differs in the groups of organisms under study. Thus distinct taxa can be recognized
5. Phylogenetic conclusions can be drawn from the taxonomic structure of a group and from character correlations, assuming some evolutionary mechanisms and
pathways
6. Taxonomy is viewed and practised as an empirical science
7. Phenetic similarity is the base of classifications
Merits of Numerical Taxonomy:
Utilizes better and more number of described characters. Data are collected from a variety of sources, such as morphology, chemistry, physiology, etc.
As numerical methods are more sensitive in delimiting taxa, the data obtained can be efficiently used in the construction of better keys and classification
systems,
Number of existing biological concepts have been reinterpreted in the light of numerical taxonomy
Numerical taxonomy allows more taxonomic work to be done by less highly skilled workers
Demerits of Numerical Taxonomy:
Useful in phenetic classifications and not phylogenetic classifications.

Proponents of “biological” species concept, may not accept the specific limits bound by these methods
If characters chosen for comparison are inadequate, the statistical methods may give less satisfactory solution
Different taxonometric procedures may yield different results. A major difficulty is to choose a procedure for the purpose and the number of characters needed
in order to obtain satisfactory results by these mechanical aids. It is necessary to ascertain whether a large number of characters would really give satisfactory
results than those using a smaller number.
ZOOGEOGRAPHICAL REALMS
Zoogeography is the sub discipline of zoology concerned with distribution of animals. It can be studied at global, regional or local scales.
First classification of zoogeographic realms by Sclater (1858) — Palearctic, Nearctic, Neotropical, African, Oriental, Australian.
Holt (2011) classified into 11 realms: 6 + Panamanian, Oceanian, Sino Japanese, Saharo Arabian, Madagascan
Largest realm — includes Europe, parts of Asia and Africa

Palearctic Sub Regions: Manchuria, Europe, Mediterranean, Siberian
Species: Llama, Red Panda, Moose, Giant Asian Salamander, Giant Panda
Sub Regions: California, Rocky Mountain, Alleghany, Canadian

Nearctic Resembles Palearctic in climate
Poor in fauna. Species: Beaver, Red Deer, Gila monster, Pronghorn
Sub regions: Chile, Brazil, Antilles, Mexico

Neotropical High habitat diversity from alpine on Andes to rainforests in Amazon basin
Region overall has high diversity and endemism: Armadillo, Tapir, Anaconda, Toucan
East, West, South African, Malagasy

African Sahara desert acts as a divide between Palearctic and African realms
Species: Gorilla, Hippo, Ostrich, African elephant
India, Indo-China, Ceylon, Indo-Malayan

Oriental Varied physical features. Includes Himalayas and rainforests of Borneo
Gharial, Orangutan, Gibbons, One Horned Rhino
Australian
Austro-Malayan, Australian, Polynesian, New Zealand
Rainforests, Deserts and Arid lands — Partly tropical and partly temperate
Fauna is unique: Marsupials, Tiger Snakes, Duck Billed Platypus, Emu
DISTRIBUTION OF ANIMALS
Distribution of animals is affected by their tolerance of environmental conditions

Principles governing animal distribution applies differently based on their habitat.
Distribution of animals is not uniform
Factors affecting animal distribution
Dispersal ability: Certain species disperse easily and have cosmopolitan distributions. Endemic species have low dispersal ability
Physical Barriers: Large bodies of water, intervening deserts, Sheer distance, salinity in case of aquatic animals, light
Mozambique Channel separates Africa from Madagascar, prohibiting dispersal of species from Madagascar to mainland Africa.
Geological Events. e.g. Glaciations, Continental drift
Disjunct distributions of animals like Tapir (South America and Malay peninsula) is likely due to populations separating during continental break down
and shift
Flightless birds also have a disjunct distribution: Ostrich (Africa), Rhea (South America), Kiwi (NZ), Cassowary (Australia)
Tolerance levels to environment: Tolerance to salinity, temperature, pressure, humidity, light etc. demarcate boundaries of species.
Composition of freshwater communities is heavily dependent on climate. Salt water organisms have a wider range because the seas of the world flow
into each other.
Similar climates have broadly similar organisms (exception being Tropical Africa and South America); Pacific Salmon has a bipolar distribution,
distributed in Arctic and Antarctic.
Eurytopic animals have wide distributions, while stenotopic animals have narrow distributions across different parameters
Competitive Exclusion: e.g. Barnacles along the tide line
Ability to establish new population in barren land
Means of dispersal
Natural rafts, driftwoods

Winds
Storms
Land bridges
Human influences
Continental Drift (Wegener)
Continental Drift refers to the process by which movements of fragments of land (continents) move on the surface of the earth.
Wegener postulated that all continents formed a single landmass (Pangea) and the seas of the world formed Panthalassa
Pangea fragmented in the Triassic (breaking into Laurasia and Gondwanaland). Consequent movements and fragmentations have led to the present day
configuration of continents
Evidences for Continental Drift
Bulge of South America fits into the bight of Africa

Identical fossils in North American and European coal deposits
Convergent mountain ranges
Rocks in Brazil and West Africa are similar
Glaciation evidences in Antarctica, Southern Africa, India and Australia
Mesosaurus fossils are found only in South Africa and South America
Terrestrial reptile fossils found in Africa and Antarctica are similar
EVOLUTION OF HORSE
Horses are placed in Order Perissodactyla, suborder Hippomorpha, superfamily Equoidea and family Equidae
Among the best known evolutionary histories, because the fossil records are remarkably complete
It offers a classical example of Orthogenesis or straight course of evolution, and one of the finest examples of progressive evolution through natural selection.
Horse fossils have been found in different parts of Europe and Asia across geological times, but the entire sequence of evolution is North America most
completely, in the sedimentary deposits, from Eocene to recent times.
General Evolutionary Trends
Increase in size and height — rabbit like animal to 6ft tall grassland animal
Lengthening of limbs, perfection of the hoof for fast running in open grasslands
Reduction of ulna and fibula, with a consequent limitation on the range of movement. Strengthening of Radius and Tibia.
Change of foot posture from semiplantigrade to unguligrade; loss of digits from five to one — Helped in faster running
Enlargement and better development of the third digit (median digit) and reduction of the other lateral digits — Perfection of the hoof
Reduction and deepening of the front portion of the skull and lower jaws to accommodate the high crowned cheek teeth.
Elongation of the preorbital or facial region of the skull and migration of eyes to the top of head.
Lengthening of the face in front of the eyes
Increase in size and complexity of brain
Reduction in pectoral girdle and disappearance of the weak clavicle.
Elongation and complexity of teeth with an increase in the height of the crown of the cheek teeth.
Pre Molars become successively Molariform
Modification of teeth from brachydont (low-crowned) to hypsodont (high crowned) - tougher food (grass).
Straightening and stiffening of the back
Nostrils became wide to allow more air into strong lungs and stamina increased
Body became streamlined, muscles tight, without loose fat, for long and sustained running.
About the size of a dog (14 inches tall, 12 lbs)

4 toes on forelimb, 3 on hind limb
Eocene
Eohippus Little or no lateral vision, short neck, short snout, short legs, long tail
60 mya
Teeth were short and crowned, for eating plants
Small brain, with small frontal lobes
24 inches tall. Longer snout, legs and neck than Eohippus

3 toes on each foot. 4th toe on forelimb was reduced to vestigial nubbin
Oligocene
Less arched back
Mesohippus 35 to 40
Legs, snout, neck and face were longer than Eohippus
mya
Larger legs enabled it to run faster
Shallow facial fossa, and a depression on the skull
24 inches tall at the shoulder, weighed more than Mesohippus

Incisor teeth form, Longer head
Overlapped with Mesohippus for about 5 mya period
Oligocene
Miohippus Facial fossa was deeper and more expanded
30 mya
Teeth changed to be better suited to chew harsh, abrasive grass
Increase in body size, leg length and length of face
Horses began to stand permanently on tip toe
Looked very similar to today’s horses

Stood over 36 inches tall at shoulder
3 toes, outside toe became weaker. Toes started turning into hooves
Teeth similar to present day horses
Miocene Head changed and eye moved, allowing better vision
Merychippus 20 to 25 Longer neck for easier grazing
mya Better sense of smell
Muzzle became elongated, jaw became deeper and the eye moved further back
Radius and Ulna of foreleg fused, so that leg rotation was eliminated — helped in
rapid running over hard ground
3 toed but spring footed
Considered direct link to Equus.

First single toed hoofed horse
Pliocene Strong leg ligaments to increase speed and power
Pliohippus
12 to 6 mya Resembled a pony
Dished face
Deep facial fossa
Does not have a dished face

Equus 5 mya No fossa (Pliohippus had a deep facial fossa)
Teeth are very straight (Pliohippus teeth are curved)
EVOLUTION OF ELEPHANT
Elephants belong to Order Proboscoidea
General Evolutionary Trends
Development of giant size

Development of pillar like limbs and short, broad and padded feet.
Lengthening of limb bones
Growth of skull to a very large size
Secondary shortening of the lower jaw
Shortening of the neck
Elongation of upper lip into muscular, mobile trunk proboscis
Development of 2nd superior incisors into tusks
Modification of cheek teeth for chewing and grinding plant food
EVOLUTION OF MAN
Trends in Evolution
Bipedalism
Led to upright posture, result of anatomical restructuring of pelvis and lower limbs.
Consistent bipedalism characterised by erect posture with straightened knees is found only in man (bent knee gait is found in some other apes as well)
Structural changes in lower limb — elongation of femur and reconstruction of foot including its digits.
Lower limb bones become larger than upper limb bones (opposite to great apes)
Foot bones and digits have shifted from ancestral grasping to a weight bearing platform
Thumb Opposability
According to Darwin, bipedalism evolved when our ancestors came to live less on trees and more on ground
Brain Size and Complexity
Rate of increase was slow initially, and increased later on
Related to tool use, tool manufacture, complex social life, ability of speech, slow rate of infant maturation which requires greater parents care.
Associated changes
Shifting of foramen magnum towards interior
Total facial plate including forehead becomes high, upright and nearly vertical
Loss of heavy eye brow ridge
Decrease of jaw length and teeth size and teeth number
Enlargement of dorsal side of skull to accommodate the larger brain
Adjustment of Behaviour
Division of labour between the sexes
Development of communication and signalling systems
Facilitated by increasing brain size
Tool Making
Extensive environmental manipulation
Facilitated by grasping hand of primates
Acquisition of meat protein
Caused changes in jaw and musculature and dentition
Hunting probably evolved in Homo erectus
Miocene (26 mya) Propliopithecus
Pliocene (3-8 mya) Dryopithecus
Australopithecus
Pleistocene
Homo erectus Neanderthal
Recent Gorilla Homo sapiens
Ape like primate, Short arms

Propliopithecus Oligocene
Ancestor of modern apes and man
Aegyptopithecus Miocene
Dryopithecus Miocene Fore limbs shorter than hind limbs
Miocene, early Ape-Man

Ramapithecus
Pliocene Found in India and Africa
Connection between man and ape

Australopithecus Pleistocene Erect posture, 4 ft tall, Bipedal locomotion, dentition like man
Hands used for non-locomotory function
Homo erectus
True Man. Also called Java man
Connection between Ape-man and Modern-man
Taller than Australopithecus
Hunter Gatherer
Homo heidelbergensis Primitive Man
Advanced primitive man

Extinct 25,000 years ago
Homo neanderthalensis
Slightly shorter than present day man. Low forehead. Large teeth
Use of tools, Speech
Extinct modern man

Cro Magnon man
Lived in caves, Hunter Gatherers
EVOLUTION OF CAMEL
Camel is a hoofed mammal belonging to the family Camelidae, of order Artiodactyla.

Now there are only two living genera Camelus (North America and Asia) and Llama (South America)
Evolutionary Trends
Gradual increase in size

Subsequent loss of lateral digits
Elongation and fusion of metapodials to form the canon bone
Change over from unguligrade to digitigrade form
Reduction in number of teeth
Gradual elongation of teeth
Phylogeny
Ancestral camels appeared during Upper Eocene or early Oligocene and went through major part of their evolutionary development in Tertiary period in North
America.
During Pleistocene, they migrated they migrated to other parts like South America and Old World because of Ice age, and became extinct in North America
Short limbed and 4 toed, the size of a rabbit.

4 primitive teeth in continuous series
Eocene Protylopus
Radius and Ulna was separate and Fibula was complete.
Narrow face, but the bony orbit was incomplete behind
Poebrotherium
Size of a sheep
Ulna was coalesced with Radius and Fibula was fused with Tibia except at the two ends.
Protomeryx
Poebrotheirum, Complete encircling of orbit by bone
Oligocene Protomeryx,
Paratylopus
General trends
Rapid reduction and loss of side toes in early camels, so that by Oligocene, the animal became bi toed.
Neck was slender with a small tapering skull
Grinding teeth of upper jaw were short crowned, while those of lower jaw had begun to elongate.
Miocene Procamelus
Pliocene Pliauchenia
Procamelus
Holocene Reduction in number of teeth and beginning of formation of Cannon bone
Llama

2A Evolution

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02/08/2019 Evolution

Features of living entities

Generation of life from non-living matter

Clay Origin: Complex organic molecules arose on a non-organic replication platform.

Prebiotic soup led to formation of short RNA molecules.

Variation: Individuals in a population have inherent variation

Essentially, NS is differential success in reproduction.

HARDY WEINBERG EQUILIBRIUM

A population in HW equilibrium is NOT evolving

P + Q = 1 — Allele Frequencies

Viability Selection: Survival until adulthood

Stabilising Selection Directional Selection Disruptive Selection

Favours a single phenotype and allele

Continuous Variations Discontinuous Variations

Variations are very small and in a graded series. Genetically quantitative

Stable and heritable changes in genetic material

Most of the mutations are inconsequential because

Mutation in various theories

At the level of an individual base

Base substitution: Transition or Transversion

At the level of chromosome

Role of Mutations in Speciation

Random, non-directional process of change in allele frequencies

Genetic Drift Natural Selection

Random and non-directional process Directional process

Bottleneck Effect Founder Effect

Geographical: Populations cannot mate due to geographical barriers.

No new populations are formed

Origin of new populations from old ones

Based on timescales (and correspondingly, divergence)

Drift + Mutation + Selection + Gene Flow = Relatively Small changes in population

Origins of new levels of biological organisation

Common features of Macro and Mega Evolution

Pre adaptations (Not necessary in micro evolution)

Monophyly: Taxa whose members have descended from a common ancestor

Based on rate of Evolution

Bradytelic Evolution: Gradualism. Slow evolution in minor ways

Mimicry provides protection to the mimic

Predator mimics a harmless model in order to gain advantage in capturing prey

Animal behaves as if they are dead.

Sudden mutation leads to phenotypic changes.

Batesian Mimicry Mullerian Mimicry

Protective mimicry in which species that is edible or harmless resembles

Factors favourable for preservation

What do fossils tell us?

Movement: Tracks of insects, trails of worms, foot prints of vertebrates

Biostratigraphic classiﬁcation of sedimentary rocks

Exceptions to Species Concept

Issues in classiﬁcation of organisms

Deﬁning species boundaries, as speciation is a continuous process.

Why are names of taxa changed?

Makes study of a wide variety of organisms easy

Holotype: Single specimen on which description of the species is based

Binomial nomenclature is the system of scientiﬁc names applied to taxonomic units

Each species is assigned a two part scientiﬁc name

International Code for Zoological Nomenclature

Ancestral Feature: Similarity derived from common ancestor of the group

Time trees using calibrations

Chromosome number and structure — active ﬁeld of study in primates

Physiology reﬂects gene expression

Can only be measured in living specimens — no fossils, no museum samples

Phylogeny can be inferred from molecular data — DNA, RNA, Protein

Molecular Taxonomy has revised many traditional notions

Protein sequences can be aligned and analysed like DNA