EVOLUTION: Lectures 1 & 2

Darwin’s argument to explain

adaptation in evolution
Robin Allaby
 Aims
• To understand the context in which Darwin constructed his theory
• To understand the basis of Darwin’s arguments and examples he used
• To understand some of the other ideas about evolution

Objectives
• To look at the history of evolutionary thought
• To deconstruct Darwin’s thesis: On the Origins of the Species
 What did people believe at the time of
Darwin?

• Immutability of species (mostly)

• Natural Theology: nature is perfectly designed with an order
and complexity that must reflect a creator.

Image of
watch to go
here

Robert Boyle Disquisition About the William Paley Natural Theology (1802)
Final Causes of Natural Things (1688)
 What is evolution and why did people not
think it occurred?

• Now evolution is understood to be organism change by descent over time

• Evolution needed a credible mechanism to explain:

complexity
the same or similar species occurring in remote parts of the world
how do we have clearly defined ‘species’ that cannot interbreed

• In order to understand why people thought what they did, one must appreciate
the Greek philosophers
 Greek Philosophy and the origins of
evolutionary thought: pre-Socratic

Thales 624-546 BCE Anaximander 610-546 Parmenides 520-450 BCE Empedocles 490-430 BCE
BCE

The pre-Socratic philosophers rejected myth and legend, and looked for explanations
of our reality in the natural world.
 Greek Philosophy and the origins of
evolutionary thought after Socrates

Democritus 460-380 BCE Plato 428 - 348 BCE Aristotle 384-322 BCE

Epicurus 341-270 BCE Zeno 344-262 BCE

 Use of the term ‘evolution’
Originally used by Bonnet in embryology to
mean ‘unfolding’, he used it to describe the
progression of beings up the chain of being

Charles Bonnet 1762

Herbert Spencer, a proponent of Natural

Selection, introduced ‘evolution’ to replace
Darwin’s ‘modification by descent’. Only
became accepted by Darwin towards the
end of this life. He also coined ‘survival of
the fittest’ that Darwin also adopted in later
editions.
Herbert Spencer
The major influences that led to the theory
of Natural Selection

William Paley
Charles Lyell

Thomas Malthus
Adam Smith
Darwin’s realizations: the main elements of the
theory of Natural Selection

Species have the capacity to The variations involved are

reproduce geometrically.
Consequently, they will have
to compete for space: there is
a struggle for existence.
Species will keep each other in
check.
tiny, but will give rise to big
differences (speciation) over
inordinately long periods of
time.

Species have variation in

individuals, therefore those
with more suitable variations
will fair better in the struggle
for existence; they will be
naturally selected.
The vast complexity of
interaction of species with
each other and environment;
the ‘watch’ is simply emergent
from the ‘selfish’ actions of
individuals – the ‘invisible
hand’.
 Other influences of the day
Georges Cuvier – father of palaeontology; immutability of species and
occurrence of extinction
Louis Agassiz – proposed recent ice age
Jean-Baptiste Lamarck – different mechanism of evolution
Alfred Russell Wallace – came to very similar theory at around the same
time

Structuralist (fomalist) vs Functionalist debate: is adaptation only a minor

factor in the morphology of organisms which sits on an archetypal
(designed) form?

Johann Wolfgang von Goethe – theories of form the leaf as the archetypal
unit of the plant
Etienne Geoffroy Saint-Hilaire – theories of form – the vertebra as the
archetypal unit of the animal
Natural Theology provided the template

‘On the Origins of

Species’ was modelled
on Paley’s arguments,
forming a counter
argument using many
of the same examples
e.g. swim bladders
and lungs.
 Darwin did not rush his argument

Voyage of the Beagle 1831-1836

On the Origins of Species by Natural Selection or the Preservation of Favoured Races in

the Struggle for Life 1859 (1st Edition)

Why did it take so long?

 Darwin’s ‘one long argument’…

1. Variation under domestication record

2. Variation under nature 10. On the geological succession of
3. Struggle for existence organic beings
4. Natural selection 11. Geographical distribution
5. Laws of Variation 12. Geographical distribution 2
6. Difficulties on theory 13. Mutual affinities of organic beings:
7. Instinct Morphology; Embryology;
8. Hybridism Rudimentary Organs
9. On the imperfection of the geological 14. Recapitulation and conclusion.

….is a 500 page abstract!!

Gould’s summary of Darwin’s argument

• Uniformity – use of small scale changes in domestication to demonstrate evolution

• Sequence – use of examples of varieties which can be linked to show a sequence of change
• Consilience – how biogeography, particularly of island biota, make sense under natural
selection

• Discordance – how seemingly nonsensical structures and species make sense under
natural selection.
Variation under Domestication
• Each domesticate type must have come from a
single wild species

• Cannot cross true breeds to produce a stable

new breed

• Crossing pigeon varieties causes reversions to

features of Rock pigeon (Darwin concluded that
to be the progenitor of all varieties) p25‡

• Breeders know they need large populations to

produce new varieties p41

• The ability to control breeding is paramount in

the production of new breeds; pigeons easy, cats
difficult p42

• Man has produced new races not by crossing old

ones, but by artificial selection of the same races

‡ pagination refers to first edition of On the Origins of Species

 Variation under Nature
‘doubtful species’
• Variation occurs in important parts of organisms as well as
unimportant; systematists had argued the opposite (and hated
variation within species which just gave them problems); it is
the awkward (intraspecific) variation that is important p45‡

• Darwin argues that defining a species is highly problematic,

many doubtful species occur which may be better viewed as
incipient species (extreme variants) p49
primrose
• Large genera have more varieties per species, and more
populous species have more variants; implies that speciation is
ongoing and that varieties are incipient species.

cowslip ‡ pagination refers to first edition of On the Origins of Species

 Struggle for existence
• the tendency for geometric expansion through reproduction for all species, even those with low
fecundity (elephants) means that without check they would over run the planet in a very low
number of generations; ergo they are in check. The check is the comes from the struggle with all
the other species as well as against the physical environment (e.g. climate). The struggle is most
intense between similar species, and ultimately individuals of the same species.

• species must exist in numbers in excess of checks; illustrates why agriculture works, but not on
the garden scale. p70

• the interrelationships of checks is highly complex; Darwin’s example of town cats affecting
numbers of red clover and heartsease around settlements.

• a lack of checks explains why alien species tend to run riot

Aliens: two examples not used by Darwin

Cane toad

Indian Balsam
 Natural Selection
[the] ‘preservation of favourable variations and the rejection of injurious variations’ (p. 81 On the Origins of
Species)

• Examples qualified by domesticates comparison

• Catskill wolf example of selection (greyhound) p90

• sexual selection (breeder’s selection of bantams) p89

• Naturum non facit saltum – comparison to Lyell’s

Uniformitarianism p95
Natural Selection

Conditions of Selection
• Good conditions for selection to occur p102:
- large population
- rapid reproduction
- locally restricted
- isolated
• predicted that birds would not evolve quickly (too much mixing of different populations), like breeders must keep
new lineages pure.

• ocean islands provide isolation resulting in a high number of endemics

• large open (continental) areas have more numbers, therefore more variants, therefore fiercer competition;
speciation is expected to be quicker in continental areas.

• Small areas/systems will have slower speciation, and where one is more likely to find archaic species (e.g.
Ornithorhyncus and Lepidosiren. Madeira’s flora reflect the extinct Tertiary flora of Europe. p107
Natural Selection

Divergence of Character
• How do small differences become big differences?
- again Darwin refers to pigeons demonstrating how traits are repeatedly selected for (e.g. beak size)
p111

• Divergence allows new niches to become occupied, allowing an increase in numbers

• Darwin singles out Australia as an example of poor diversification – marsupials everywhere but insubstantially
different, so fail against European immigrants; concept of phylogenetic advantage

• Ancestral forms quickly driven to extinction; Darwin recognized the corollary for extinction that very few
ancestral species could have given rise to extant ones
 Laws of Variation
• what is real heritable variation, and what is ‘conditions of life’? E.g. fur
thickening in colder climates - Darwin thought adaptation to climate
overstated – domesticated animals move easily between biomes p139

• effects of disuse e.g. Madeira’s wingless beetles p134

• correlation of growth e.g. deaf blue eyed cats p143

• law of compensation ‘in order to spend on one side, nature is forced to

economize on the other’ Goethe e.g. cows milk/fat Brassica foliage/oil
p147

• traits specific developed in one species will tend to show greater

variation then the rest of the genus , because NS is still in action p150

• analogous variation will occur between closely related species, i.e.

they are predisposed to do similar things (independently) e.g. swedes
and turnips. p159
 Difficulties on Theory
• Why do we not see innumerable transitional forms – i.e. why are all species not joined up?

• How can you have large transitions in habit?

• Can instincts be acquired through Natural Selection?

• Why is it that species being crossed are infertile, but varieties when crossed are fertile?
Difficulties on theory

1. Why are species discrete?

• intermediate forms are outcompeted

• more of a problem with hybrid zones – Darwin cites another sheep

example p173

conditions best
for intermediate
(hilly track sheep)

Intermediate zone only

small, therefore
cannot produce
variants as quickly as
either A or B, therefore
no intermediate
variant becomes
established from the
intermediate
conditions best conditions best population – they
for variant A for variant B come from A or B
(plains sheep) (mountain
sheep)
Difficulties on theory

2. Large transitions in habit

• Darwin saw the transition from land to water as fairly minor to explain
e.g. Mustela vison (Neovison vison), American mink – webbed feet with fish
eating habit in summer, and polecat habit during the winter; an example of
a transitional species. p179
• But flight, and bats in particular, he considered a real problem – no
transitional allied species. He cites squirrels/flying squirrels, and
lemurs/flying lemurs as possible examples of the nascent evolution of
flight. Birds wings also have many uses other than flight. Problem is that
intermediate forms would be outcompeted and extinct.
• Organs of extreme perfection; the eye the most difficult to explain.
Darwin uses examples of redundancy of function as a possible mechanism:
swim bladders evolving to lungs (although they didn’t), taking one of
Paley’s examples from Natural Theology. p186
 A bit more on perfection
No organ will be formed for the purpose of causing pain or for doing an injury to its
possessor. William Paley

Darwin argued that the degree of perfection was only in

relation to the other inhabitants in the immediate struggle:

New Zealand inhabitants – well adapted to each other, but

hopeless in the face of European invasion; the organisms of
smaller islands/areas will yield to those of larger ones. P.201

Imperfection of bee sting: modified boring instrument which

has backward serrations, the poison originally adapted for
causing galls. p202
 Instinct
Instincts evolve just like organs, Darwin argued
• examples from domestication:
- behaviours associated with breeds of dog; pointers (point), retrievers
(retrieve), and sheepdogs run around sheep rather than at them.
- chickens have danger chuckle that causes chicks to hide in grass (a
useless trait now). p216
• cuckoos – behavioural adaptation to staggered egg laying habit p217

• Slave making ants – evolved from stealing pupae for food, to rearing for
slavehood. Formica sangiunea is half-way dependent, Polyergus rufescens is
completely dependent on slaves. p219
• Hive bees wax comb – perfect economization of wax, mathematically. A
sequence of increasing complexity made by simple actions can be seen across
species. p224

‘Humble’ bee Melipona bee Honey bee

 A bit more on bees
Bees provide an apparent problem for Natural Selection
that worried Darwin greatly p236:

sterile workers do not leave offspring, how did they

obtain a difference in structure to their parents?

Darwin argued that selection acts on the family level,

using domestication again as an illustration;

Farmers slaughter beef from good stock in the hope that it

will yield good meat, ie they select on the level of the
family.‡

Darwin cites this as proof against Lamarckism (inheritance

of acquired characters), because no amount of experience
of the neuters could have affected their evolution, since
they do not reproduce. P242

‡ Note that this group selection is not how we view bees

now, but the sterile workers as effectively an extension of
the individual on whom selection is acting (the queen).
 Hybridism
• considers two categories of sterility p246:
- when species are first crossed (very infertile)
- sterility of hybrids (become more fertile with generations)

• experiments of the day showed that some species were more fertile when crossed
than bred intraspecifically p250

• animals generally less interfertile, although cites taurine and zebu cattle; thought
that domesticates generally were hybrids that became more fertile p252
• fundamental message: there is no line at which varieties end and species start, all
one continuum; therefore species ARE varieties, varieties are incipient species and
sterility of species is not a ‘designed’ characteristic. p263
Hippeastrum sp.

Passiflora sp. Verbascum sp.

 The Imperfections of the Geological Record
The geological record was a big problem for Darwin; not many good sequences of evolutionary change (no
examples in Origins), but plenty of large jumps and extinctions. Darwin criticized the geological record as a
source of evidence because of the following points:

• fossils are only laid down under particular, and quite sporadic circumstances; mostly when seas are
shallow and the rate of sedimentation is rapid and equaled by subsidence of the ground (ie land is
sinking) p291

• land loss is a time of extinction, unlikely to find new forms appearing. Periods of elevation are not
associated with fossil formation. p292
• immigration and emigration in fossil record can look like extinction or sudden speciation. p294

• palaeontologists tend to over ascribe species p297

• new forms are likely to be highly fecund and locally restricted; unlikely to observe transitional
stages in geological record. Sudden appearance of species likely to be spread of these ‘established’
new forms. Examples of the day include monkeys whales and cirripedes p298
• Darwin noted that oldest fossils of the day (Silurian) were already diverse in form (trilobites,
nautili), therefore much older fossiliferous strata must have existed, or exist somewhere. He was
right! p306
 Geological Succession of Organic Beings
Darwin ‘needed’ the geological record to demonstrate the mutability of species. The most striking thing
about this chapter is the dearth of actual fossil examples!
• Catastrophism generally given up amongst geologists at this time; species should gradually go
extinct one by one, although he accepted that sudden extinction episodes did occur (e.g. end of
‘Secondary’ period ammonites vanished, and end of the Palaeozoic period trilobites vanished). p317
• Forms of life changed simultaneously throughout the world. New World forms resemble European
fossils more than extant European forms. P322
• Fossils can all be placed in relation to extants forms either within or between groups, older fossils
more different. Fossils connect ancient lineages to the rest of life – e.g. Lepidosirens
• Fossils should approach one another in similarity away from extant forms, Darwin claims this to be
true. p333
• Intermediates should occur in mid succession, Darwin claims this to be true of Devonian forms
between Silurian and Carboniferous (but elephant group presents problems). P334
• Later forms of fossils should be more highly developed. p336

• Similar fossil and extant forms occur in same areas e.g. armadillos in La Plata (South America), and
rattites in New Zealand, marsupials in Australia. p338
 Geographical Distribution
• similar climates around the world have “utterly” different faunas, but within regions with different
climates similar fauna occur. P347

• barriers correspond to abrupt fauna changes e.g. Panama isthmus, as compared to the East African
coast and the Pacific islands.
• niches are filled in parallel in different areas by different species allied to their region (e.g.
Vishcacha/rabbit.

• Disjunct species distributions explained by migration (alternative of the day was independent
creation) – why then have different species in Australia, S America and Africa where conditions are
the same?
 Means of dispersal
• climate can open close migration routes, sea level changes

• islands more of a problem:

- not recently connected to continents (rocks all young) p356
- continents too distinct in fauna to have been recently connected
- Darwin’s seawater experiments with seeds – many species survive well after a long time in
the water (e.g. 64/87 seeds of various species germinated after 28 days) p 358
- birds efficient dispersers of seeds
- icebergs carry soil; e.g. the flora of Azores is northern (also has erratic granite rocks) from
ice flow. p363

• glacial periods recognized as very important (had recently been put forward by Agassiz in 1837)
- alpine forms trapped up mountains (islands) were perturbed both on warming and cooling
therefore expect to see many ‘doubtful forms’ and varieties, which Darwin claimed to be true.
- Darwin believed that many northern hemisphere species crossed the equator during the last
ice age to explain their presence in the southern hemisphere. p369
Geographic distribution continued
• fewer species on oceanic islands p388

• but more endemics – explained by isolation (islands on migration

routes do NOT have many endemics e.g. Bermuda) p390
• poor competitors relative to continental invaders

• remote islands deficient in classes – niches filled by atypical groups

• Batrachians almost never found despite good conditions (introduced

to Madeira, Azores and Mauritius where no a nuisance)
• mammals generally not found more than 300 miles from continental
mainland (except bats)

• Galapagos islands have 25 endemic bird species, all resemble

American species despite very different conditions there, relative to
Cape de Verde (which resembles African fauna)

• why do species not spread through islands, but tend to be locally

endemic? Local adaptation e.g. Galapagos mockingbird means each
species is locally ‘fitter’ than the others (note – not separate niches).
p402
 Classification: morphology
Darwin argued that modification by descent naturally lends itself to
classification. Previous classifications were thought to reveal the ‘plan of the
creator’, but Darwin thought that Linnaeus’s classification partially revealed
evolutionary history. p411
• at this time features that are external and adaptive were considered important for classification –
Darwin argued that the opposite is true, should rely on features not unduly influenced by the ‘conditions
of life’. Reproductive structures good, embryonic structures very strong. p414
• adaptive resemblences give false unity – agreed with Lamarck (who was first to point it out) on this
point e.g. fins of fish and whales. P427
• Aberrant species (Ornithorhyncus and Lepidosiren) sit on the end of many extinct intermediates; lucky
members of orders that have otherwise been outcompeted p429
• Members of one order may resemble another – but only generally, not specifically e.g. Viscacha (rodent)
has some marsupial-like features, phascolymys is a marsupial that has some rodent-like features p430.

• use ‘unity of type’ – features which are homologous between species e.g. pentadactyl limb, and within
organism e.g. legs and mouthparts of crustaceans, or skull and vertebra. Darwin used these relationships
as more evidence against a creationist explanation – they make sense under Natural Selection
 Classification: embryology
Darwin argued that ancestral forms are present at embryonic stages: ‘the
embryo is the animal in its less modified state, and in so far it reveals the
structure of its progenitor’ p449
• Embryonic features are generally not subject to the conditions of life:
selection does not act on them, and so does not actively remove them. E.g.
chicken teeth, provides link to archosaurian ancestry

• The phylogenetic placement of an organism may be easier from the embryo

than the adult, because retention of ancestral features bridges the gap that
would have represented by intermediate forms, e.g. cirripedes assigned to
crustaceans

• Darwin again resorts to domesticate forms to illustrate:

Archosaurian teeth in chicken embryo - bulldog and greyhound puppies differ less than adults
- race horse/ cart horse young
- young pigeons almost indistinguishable
- all because breeders select for features in the adult

‘ancient and extinct forms of life should resemble the embryos of their
descendents ’ p449
 Classification: rudimentary organs
‘rudimentary organs may be compared with the letters in a word, still retained
in the spelling, but become useless in pronunciation, but which serve as a clue
in seeking for its derivation’ p455

Examples of rudimentary organs given by Darwin:

• mammae in male mammals

• bastard-wing (alula) of birds

• rudimentary lung-lobe in snakes

• rudimentary pelvis in some snakes

• Teeth in
foetal wales (upper jaw)
upper jaw of unborn cows
beaks of some birds
• insect wings soldered shut

Rudimentary organs tend to be larger in embryos because of weakness of selection at that stage

The existence of rudimentary organs is useless, imperfect and a waste; nature is not always beautifully
designed (as per Natural Theology); they only make sense under the theory of Natural Selection.
 Summary
Difficulties (where he expected Summary of arguments
to be criticized)
• variation evident in domesticates markings or horse stripes)
• origin of complex organs
• struggle for existence self-evident • NS explains why features which define a
• The sterility of species species tend to be more variable than
• varieties explained as incipient species rest of genus
• the dispersed location of allied
• large genera will have species with • NS explains instinct evolution in small
species around the world
many varieties steps
• the lack of intermediate forms
• distinction of species due to new • geological record imperfect, but still
varieties outcompeting ancestral forms supports descent with modification
• not all varieties and species can survive • NS explains biogeography

• adaptation is only driven as far as • NS explains the peculiarities of islands

competition requires – can be inept and
wasteful
• NS explains disuse of organs
• NS explains reversions (pigeon
• all life falls under one scheme, in which
adaptive characters are less useful than
embryological and rudimentary ones.

Darwin’s Entangled Bank

‘It is interesting to contemplate an entangled bank, clothes with many plants of many kinds, with birds singing on the
bushes, with various insects flitting about, and with worms crawling through the damp earth, and to reflect that these
elaborately constructed forms, so different from each other, and dependent on each other in so complex a manner, have
all been produced by the laws acting around us.’ p489