SPECIES CONCEPT

INTRODUCTION

An historical review of species concepts reveals the great difficulty and controversy biologists have
experienced in attempts to develop a species concept of general application. As a result of the many complex
ways in which evolution plays out across the many different types of biological organisms, we may not arrive
at a general, unified concept.

Species concepts have focused on three major aspects of species: 1) morphological characteristics (phenetic
characteristics = phenotypic, especially mathematically quantifiable ones) used to distinguish species, 2)
biological properties which keep species separate (reproductive isolation), and 3) biological properties which
maintain species (fertilization and genetic cohesion). But these are not the only issues involved with trying to
develop a concept of species.

In considering species concepts, one may consider whether each concept theoretically identifies natural units
and whether the discrete clusters of phenetic traits we use to identify species are explained by the concepts.
Moreover, some concepts define species at one instant in time whereas others attempt to define species
through geological time. Some address the process of speciation whereas others address the products of
speciation.

Experiments are not effective in solving the species problem, and studies based upon the comparative
method and a dialectic approach -- intellectual inquiry involving reasoning through dialogue among scientists -
- have been the main means of studying the many varied life forms on earth.

Historically, our concept of a species has changed and is still changing. Six major concepts have emerged from
the pages of the biological literature. [Note: The textbook (BE: p. 60) also discusses Greek species concept,
tyological species concept, and Darwin's species concept. Others concepts also exist, e.g. Van Valen's
(1976) ecological species concept and the phenetic species concept.]

1. Morphological Species Concept

2. Biological Species Concept
3. Evolutionary Species Concept
4. Recognition Species Concept
5. Cohesion Species Concept
6. Phylogenetic Species Concept

MORPHOLOGICAL SPECIES CONCEPT

Morphological similarity (or dissimilarity) is the sole criterion for determining species. Thus, the degree of
individual morphological similarity or difference (vs. distinctiveness) is the primary, decisive criterion of species
status. This concept involves subjectivity in the definition through the degree of difference. Different species
are organisms that look different.

Species identified under this concept and soley on the basis of morphological differences are
called morphological species or morphospecies. This concept is fundamentally an application of
the typological species concept.
Some critics of the phenetic species concept view it as a numerical update of the morphological species
concept.

Some problems with morphological concept include: 1) sibling species, 2) sexual dimorphism,
3) polymorphisms.

Sexual dimorphism
Different morphological forms occur in the two sexes.

Example: Males and females of mallard and pintail ducks are distinctly different.

Polymorphism
Two or more morphological forms in a population. [Polytypic (many types) is term often referring to variations
among populations; may be used in reference to any taxon with 2+ subtypes.]

The morphological species concept overlooks the secondary role of morphological differences in species
formation. Morphological differences among species are a secondary by-product of genetic divergence -- not
the cause of it!!!

Example: peppered moth Biston betularia

Sibling species
Species which are exceedingly similar or nearly identical morphologically and have sympatric populations
which are reproductively isolated from one another. [Sympatry means living in the same place and refers to
populations of two or more species living together in the same area.]

Often sibling species are first detected through studies of behavior or mating calls. The study of non-
morphological traits shows that organisms once thought to belong to one species actually represent two or
more species whose small morphological differences previously were unrecognized.

Examples: Sibling species have been found based on mating calls in crickets and tree frogs.

This illustrates the fact that using the degree of morphological difference to define species is subjective, just
as are cases involving great morphological differences in forms that freely interbreed.

BIOLOGICAL SPECIES CONCEPT

Dominant concept in evolutionary literature today, at least among zoologists.

Mayr (1940) defined species as "groups of actually or potentially interbreeding natural populations which
are reproductively isolated from other such groups." Dobzhansky (1950) defined species as "the largest and
most inclusive ... reproductive community of sexual and cross-fertilizing individuals which share a common
gene pool."

The most defining aspects of this concept are interbreeding among conspecific populations and
reproductive isolation from non conspecifc populations. Morphological distinctness is not a criterion.
One major emphasis is that coexisting populations of separate "species" do not interbreed (reproductive
gap between "good" species; non-dimensional concept). Thus, they are reproductively isolated from each
other by reproductive isolating mechanisms (RIMs). Isolating mechanisms are a variety of means by which
intraspecific reproduction is insured. They act by inhibiting different species from interbreeding successfully.
Hence, RIMs define the limits of the gene pool of a species.

The second major emphasis is that species are viewed as representing the total collection of gene pools in all
the demes of individual species (unlimited gene exchange among populations within a
species; multidimensional concept).

Six tenets of biological species concept:

1. Species consist of populations, not unconnected individuals.

2. Species are defined by the reproductive isolation of populations, not by the fertility of individuals.
3. A species is a reproductive community.

Members of one animal species respond to one another as potential mates, and seek one another for
the purpose of reproduction.
4. A species is an ecological unit.

Regardless of the type of individuals composing a species, it interacts as a unit with other species, with
which it shares the environment.
5. A species is a genetic unit.

A species consists of a large, intercommunicating gene pool, and individuals are temporary holding
vessels of a portion of the gene pool. A species is a community of gene pools in demes of each
individual species. This aspect allows integration of population genetics into questions regarding
speciation.
6. Species are determined on the basis of distinctiveness rather degree of difference.

EVOLUTIONARY SPECIES CONCEPT

Developed because biological species concept could not be applied to asexual organisms or temporal
sequences of species where species changed over geological time.

Simpson's (1951,1961) version is addressed in BE, p. 63. This was orginally proposed to address
palaeontological sequences of species in which one species changed into another over geological time (A->B-
>C->D), which is called phyletic evolution. Today phylogeneticists regard speciation as a (branching)
dichotomons event.

Grant's version (1971) [see BE, p. 63] was developed from Simpson's to address concerns about biological
species concept.

Most recently this concept was espoused by Wiley (1978, 1981) in a form modified from earlier authors,
especially Simpson (1951, 1961). It was modified to address earlier criticisms and to resurrect it.
An evolutionary species "is a single lineage of ancestor-decendant populations of organisms which maintains
its identity from other such lineages [in space and time] and which has its own evolutionary tendencies and
historical fate".
This concept avoids problems of biological species concept by being compatible with a greater range of
reproductive modes. The biological species concept applies only to species with sexual, biparental
reproduction; asexual reproductive modes pose a problem for biological concept because, in some cases,
species are a number of clones, each clone being individuals from one progenitor.

This concept is applicable to extinct organisms, and it is compatible with various modes of speciation.

The concept has been asserted to be, "the best bridge principle between evolutionary process[es] and
pattern[s] of descent." It, however, is not a mechanisticdefinition as it addresses only the products of
evolution.

RECOGNITION SPECIES CONCEPT Patterson (1985)

Patterson's (1985) definition of a species is: "... that most inclusive population of individual biparental
organisms which share a common fertilization system."

This concept focuses on those biological traits (reproductive traits) involved with the reproductive systems
within species: fertilization processes and genetic compatibilities.

Highly controversial, but has had limited support.

Only applicable to sexual, biparental organisms.

Species are considered to be incidental consequences in the evolution of sexual reproduction. New species
arise as a fertilization system undergoes adaptation to a new habitat to ensure effective fertilization.

Three tenets of recognition species concept

1. The individual organisms comprising a species share a specific mate recognition system (SMRS) to
ensure effective syngamy within a population of organisms occupying their preferred habitat.
2. The biological traits composing the SMRS are adapted to function effectively in the preferred habitat.
3. A new species arises when all members of a small, isolated subpopulation of a parental species have
acquired a new SMRS. This facilitates syngamy under the new conditions and makes effective signaling
impossible between parental and daughter populations.

Specific Mate Recognition Systems (SMRS)

Signal-response interactions, including pollen/stigma interactions and sperm/ova interactions.

Fertilization processes that result in conspecific mate recognition includes: courtship displays, reproductive
timing, habitat selection, coloration, endocrine system control, copulatory organs, gametic compatibility, and
genetic compatibility. All above traits insure successful production of offspring. These are processes
which maintain a species as a biological entity. They not only ensure mating, but prevent hybridization.
This concept contrasts with biological species concept, which focuses on isolating mechanisms hence the
phenomena which isolate species or lead to isolation during speciation. The recognition species concept views
isolating mechanisms from different perspective, e.g. it considers courtship rituals to be facilitators of
reproduction (vs. isolating mechanisms). Under this concept, isolation function may arise as by-product of
evolution of other functions, but is not part of speciation process.

Patterson (1985) renamed biological species concept the isolation species concept, to differentiate it from his
own ideas. He called his concept the recognition species concept. Isolation and recognition are, however, two
sides of same coin.

COHESION SPECIES CONCEPT Templeton (1989)

Templeton's (1989) definition of a species is: "... the most inclusive population of individuals having the
potential for phenotypic cohesion through intrinsic cohesion mechanisms."

Cohesion mechanisms (modified from Templeton, 1989) include but are not limited to:

1. Mechanisms that promote genetic cohesion (identity) and thereby determine the limits to the spread
of new genetic variants through gene flow withinspecies.
a. Fertilization systems -- allows organisms to exchange gametes leading to successful
fertilization.
b. Developmental systems -- allows products of fertilization to give rise to viable and fertile
adults.
2. Mechanisms preserving genetic cohesion (identity) through the lack of gene flow with other
species: reproductive isolating mechanisms.
3. Mechanisms that define the fundamental niche of a species and the limits to the spread of new
genetic variants through genetic drift and natural selection.

Genetic drift promotes genetic cohesion (identity) through descent from a common ancestor.

Natural selection promotes genetic cohesion (identity) by favoring the fixation of a genetic variant.

Adaptive transitions (adaptive changes) are constrained by:

Mutational constraints that limit origin of heritable phenotypic variation

Limits (constraints) on the fate of heritable variation which include ecological constraints,
developmental constraints, historical constraints, population genetic constraints

Templeton argued that isolation was irrelevant function, rather a by-product, in speciation process because it
is a negative phenomenon and could not be favored by natural selection. He focused on what he thought was
more important: mechanisms that function prior to mating.

Explains mechanisms responsible for speciation and their genetic consequences by incorporating population
genetics.
Focuses on mechanisms that maintain genetic and phenotypic cohesion in those groups of populations we
recognize as belonging to different species, as does the biological species concept. Thus, it focuses on those
mechanisms that result in and maintain the genetic and phenotypic cohesion of species. These are
mechanisms that drive the evolutionary process, resulting in speciation.

Max King -- says this is essentially rewritten BSC (=biological species concept) and he's right.

Applicable to bisexual and asexual species because species are defined in terms of genetic and phenotypic
cohesion, as with evolutionary species concept..

PHYLOGENETIC SPECIES CONCEPT CRACRAFT (1983, 1989)

Species are defined as "... an irreducible (basal) cluster of organisms, diagnosably distinct from other such
clusters, and within which there is a parental pattern of ancestry and descent." In other words, 'A species is
the smallest diagnosable cluster of individual organisms within which there is a parental pattern of ancestors
and descent' (Cracraft, 1983).

Cracraft responded to the perceived failure of the BSC to resolve the process and pattern of taxonomic
differentiation and to the emphasis of BSC on reproductive isolation. He thought that if reproductive isolation
was not considered the critical aspect of taxonomic differentiation, that the differentiation was the primary
occurrence and interbreeding was secondary.

Cracraft argued that species should be defined on the basis of the products of evolution, rather than the
processes that produce those products. It attempts to define species throughout their evolutionary history.

This concept focuses on biological traits used to differentiate one lineage of organisms from another and the
branching patterns, hence, the nodes where lineages diverge. These are aspects of interest to systematists;
however, this concept suffers from deficiencies stemming from not addressing the evolutionary processes that
result in the branching.

This concept is best applied in the analysis of evolutionary lineages involving ancestor-descendant
relationships and the characteristics used to generate phylogenies: cladograms, branching patterns showing
evolutionary (ancestor-descendant) relationships; synapomorphies, traits shown by two or more taxa which
also was shown by their most recent commom ancestor in which it also was shown.

SPECIATION
INTRODUCTION

Individual organisms of one species usually differ from individuals of other species in one or more
characteristics involving behavior, ecology, or morphology. Species usually also differ genetically from one
another, and most, if not all, species are reproductively isolated from each other.

The critical event in the origin of a new species is the establisment of reproductive isolation between
speciating populations. Thus, we need 1) to know what kinds of barriers to interbreeding occur and 2) to
understand how reproductive isolation may arise between a parental species and a newly evolving species.
These are our main objectives. Before we consider these two aspects of speciation, we need a little
background information.

Geographic Relationships of Evolving Species

A newly evolving species theoretically can have one of three geographical relationships with its ancestral
species. These are:

1. allopatric -- the new species evolves in a separate geographic area,

2. parapatric -- the new species evolves in a contiguous geographic area,
3. sympatric -- the new species evolves in a geographic area that is included in the range of its ancestor.

Geographic Variation

Geographic variation in biological characteristics occurs in all species. Not every characteristic of a species will
vary, but we can expect variation in some traits. Geographic variation extends beyond visibile characters to
gene frequencies. Mayr (1963, Animal Species and Evolution, Chapter 11) concluded the following:

• "every population of every species differs from all others"

• "the degree of difference between different populations of a species ranges from almost complete
identity to distinctness almost of species level"
• "all characters employed to distinguish species from each other are known also to be subject to
geographic variation"

Thus, we can assert that geographic variation is nearly ubiquitous among organisms, occurring in most species
of plants and animals. Exceptions include highly vagile species such as migratory birds. Moreover, geographic
variation is seemingly pervasive, affecting nearly any specific characteristic: morphology, size, color pattern,
behavior, physiology, karyotype, gene frequencies.

Studies of geographic variation began with Charles Darwin and Alfred Russel Wallace. For example, Darwin's
observations of different island populations in the Galapagos Islands led to development of the Theory of
Natural Selection. Moreover, the study of geographic variation also was important in the development of
the Modern Synthesis in the early 1900's as debate over the genetic and evolutionary significance of
geographic variation played a key role.

The study of geographic variation has been one of the most important approaches to the study of evolution.
Evolutionary biologists have used the varying levels of differentiation among populations and species
to infer the course of evolutionary change. For example, the nature of variation indicates the nature of
adaptationsand the constraints upon them. Observations of geographic variation have indicated
that evolution is generally a gradual process because differences among populations range from those which
are immeasurably small through varying degrees of differentiation up to differences characteristic of different
species. The study of geographic variation has been very important in understanding processes of species
formation (speciation) as species showing great degree of genetic variation are more likely to undergo
speciation process.
Geographic variation in individual biolgical traits within a species may be discontinuous (distinct)
or continuous. An example of continuous geographic variation is a cline. A cline is continuous gradient of
gradual change in a trait (e.g. body size) among populations of a species over a geographic range along a
geographical transect. It is a character gradient along a continuum between extreme character states/values.
An example is body size of white-tailed deer (Odocoileus virginianus), which increases gradually with
increasing latitude in North America. The pattern probably exists because natural selection favors larger body
size at higher latitudes and the deer populations are adapted to their local environments.

The causes of clines have been debated for a long time. Most geographic variation is probably, adaptive,
hence, attributable to natural selection. Overall, factors that may be important in establishing clines include:

1. Geographic variation in forces of natural selection pressures resulting in locally adapted populations.
2. Interbreeding between formerly isolated populations that have undergone secondary contact (contact
between formerly isolated populations that diverged from a common, ancestral stock). The area of
interbreeding is called the hybrid zone.
3. Genetic drift which is not adaptive!
4. Phenotypic plasticity which is non-heritable variation.

REPRODUCTIVE ISOLATING MECHANISMS (RIMs)

Isolating Mechanisms

"Biological properties of individuals that prevent the interbreeding of populations [of different species] that
are actively or potentially sympatric." [Speciation = "reproductive closure" (Wake and Yanev, 1986)]

Refers to internal/inherent properties of species populations. Excludes external factors such as geographic
barriers. Isolating mechanisms are properties of species that serve to safeguard reproductive isolation.

Isolating mechanisms have a partial genetic basis. The evolution of new species represents the evolution of
genetic barriers to gene flow between populations through isolating mechanisms.

Isolating mechanisms often do not operate individually as absolute barriers to interspecific matings. Usually
a number of mechanisms operate as partial barriers which reinforce each other, resulting in almost total
reproductive isolation.

Reproductive isolation does not mean that two species are absolutely prevented from crossing with each
other. Species occasionally do cross in nature. Reproductive isolation means that the majority of the members
of a species habitually do not cross with other species.

Reproductive isolation and sterility are not synonymous terms. Many distinct, reproductively isolated
sympatric species are known which are not isolated from each other by sterility barriers.

Example: Anas platyrhynchos, mallard duck, and Anas acuta, pintail duck.

Adults are fully interfertile in captivity, as adults may be crossed without any apparent reduction in fertility,
and F1, F2, and F3 hybrids are fully fertile.
Based on such information, one might expect complete interbreeding in nature, where the two species are
sympatric in eastern North America.

Contrary to this possibility, few hybrids are produced!

Members of each species preferentially mate with their own species.

Hybrids show aspects of mating rituals of both species and are not selected as mates by either species.
Factors other than sterility are operating to keep these two species apart!

Types of Isolating Mechanisms

Premating Mechanisms

Mechanisms which act prior to mating to prevent interspecific crosses.

The potential waste of gametes and reproductive effort (both costly!) on unsuccessful attempts at
reproduction is not as great when premating mechanisms are operating, as compared with postmating
mechanisms. Moreover, hybrids formed during interspecific matings may use up ecological resources. Thus,
premating mechanisms are highly susceptible to improvement by natural selection.

Postmating Mechanisms

Mechanisms acting after matings which reduce the success of interspecific matings once they have occurred
by reducing the chance of fertilization and the viability or fertility of hybrids after zygote formation. Thus,
they prevent fully successful hybridization from occurring.

They do not prevent wastage of gametes or reproductive effort as effectively as premating mechanisms.
Moreover, ecological resources may be wasted by the hybrids and their progenies.

Improvement by natural selection is indirect because these mechanisms develop as by-products of genetic
divergence as populations adapt to different environments. As populations diverge genetically, genes are less
likely to interact harmoniously in a hybrid.
[Note: Some authors divide RIMs into prezygotic and postzygotic mechanisms.

Overview of Isolating Mechanisms

I. Premating Mechanisms

{Prezygotic mechanisms}

I.A. Potential mates do not meet, at least not in reproductive condition

1. Habitat Segregation or Ecological Isolation

Potential mates live in different habitats to which they are specifically adapted for survival and
reproduction. Habitat segregation may be very slight in terms of distance.
 Example: Quercus coccinea (scarlet oak) prefers wet, poorly drained soils and Quercus
velutina (black oak) prefers drier, well-drained soils. The two species of oaks are adapted to
different soil types. Hybridization occurs in intermediate man-made, disturbed habitats. Thus,
habitat isolation appears to be important.
2. Temporal and Seasonal Isolation

Different species populations are reproductively active (mating or flowering) at different times
of the day (temporal) or during different seasons.

Example: In toads, Bufo americanus breeds earlier in spring than Bufo fowleri, but there is some
overlap which is reinforced by habitat segregation. Again, hybridization occurs in man-made
habitats.

I.B. Potential mates meet but do not mate

3. Ethological Isolation

Incompatibilities in mating behaviors are barriers to mating. Thus, sexual attraction

between females and males is absent or weak.

Involves species or mate recognition through cues such as visual stimuli (plumage,
courtship displays), auditory stimuli (calls), and chemical stimuli (phermones).

Example: Ducks (Anas), both the mallard and pintail.

I.C. Copulation (or pollen transfer) attempted but no transfer of sperm occurs

4. Mechanical Isolation

Size and shape incompatibilities impede sperm and pollen transfer by forestalling it.

Animals: particularly important in arthropods (insects and crayfish); genitalia or genital

armatures act in lock-and-key fashion.

Plants: flowers have specific structures for pollination by particular pollen vectors; locations are
specific for both plant and pollen vector.

II. Postmating Mechanisms

II.A. Sperm transfer takes place but egg(s) is(are) not fertilized

5. Gametic Mortality (Gametic Isolation, if prezygotic)

Gametes are not attracted to each other, are destroyed or die before fertilization, or sperm are
incapable of penetrating egg membrane and die.

Examples:
 Drosophila -- "insemination reaction" -- swelling of walls of reproductive tract in females results
in death of sperm; sperm encounter antigenic reaction.

Sea urchins -- male and female gametes from different species do not react to each other due
to protein receptors on eggs which are sensitive only to proteins of sperm from the same
species.

Plants -- pollen may fail to form a pollen tube.

{Postzygotic Mechanisms}
II.B. Egg is fertilized but zygote or embryo dies

6. Zygotic mortality

Zygote dies, probably from an inability of chromosomes to pair during first mitotic event.

Example: occurs in some leopard frogs in Rana pipens complex.

7. Embryonic or Larval Mortality

Embryo or larva dies during development. Development of a hybrid egg is often irregular and may stop
at any stage. Involves nuclear-cytoplasmic activity as nuclear gene activity begins to be expressed.

Example: In crosses between Rana pipiens and Rana sylvatica, the hybrids do not develop beyond early
gastrula stage.

II.C. Zygote produces F1 hybrid of reduced viability

8. Hybrid Inviability

Hybrid inferiority: involves the death of a hybrid any time before maturity; also includes the inability to
produce offspring for any reason other than sterility. Ecologically they are less well adjusted to
available niches. Ethologically they are less successful in courtship. Jointly, they reduce chances of
leaving offspring.

Example: Buttercups & their hybrids. Rannunculus millanii occupies wet habitats and Rannunculus
dissectifolius occupies dry habitats. Their hybrids cannot compete with parental species in parent's
habitat; hybrids occupy intermediate, disturbed habitats.

II.D. F1 hybrid is fully viable, but partially or completely sterile

9. Hybrid Sterility

Very important in plants.

Inability of hybrids to produce normal number of viable gametes.

May be complete or partial effect.

 Not necessarily correlated with viability as sterile hybrids often show hybrid vigor. For example: the
mule (horse Equus caballusx donkey Equus asinus) shows hybrid vigor but is sterile. Hybrid vigor does
not constitute Darwinian fitness.

II.E. Viable, fertile F1 hybrid produces deficient F2

10. Hybrid Breakdown (F2 Breakdown)

Sometimes considered an aspect of hybrid sterility.

Viability or fertility of descendants of F1 hybrid is reduced.

Recombination of parental genes in hybrid gives rise to genotypes of reduced fitness.

Also occurs when there is a backcross to parental species.

Example: Drosophila pseudoobscura and Drosophila persimilis may be hybridized in the laboratory. The
F1 hybrids are fully vigorous, and although the males are sterile, the females are completely fertile. In
backcrosses involving F1 hybrid females with males of either parental species, the offspring have
reduced viability and fertility.

Summary Example of Isolating Mechanisms

Dobzhansky (1951): Drosophila pseudoobscura and D. persimilis are sibling species which have completely
isolated gene pools in nature. The are separated by four isolating mechanisms, two of which are premating
mechanisms and two are postmating mechanisms.

Premating mechanisms are:

1. Habitat -- preferred temperatures and humidities

2. Temporal -- time of day when mating occurs

Postmating mechanisms are:

3. Sterility of hybrid males

4. Hybrid breakdown in offspring of F1 female hybrids

MECHANISMS OF SPECIATION

Species are reproductively isolated groups of organisms. The question of how speciation occurs is equivalent
to the question of how reproductive isolation arises between groups of populations because reproductive
isolation is the crucial process. Issues include the following questions. Does speciation require geographic
separation? Does complete reproductive isolation evolve during geographic separation, or does isolation begin
during separation with it being completed only when an incipient species comes into contact with with its
ancestral population?
 Three major aspects of speciation process:

1. Spatial Relationships (already discussed)

a. Allopatric
b. Parapatric
c. Sympatric
2. Tempo
. Gradual: populations; many generations
a. Abrupt (saltational, quantum, instantaneous): individuals (one offspring or offspring of one
mating); one-few generations
3. Genetic
. Genic
a. Chromosomal

Allopatric or Geographic Speciation (Examples in BE, p. 75-76)

A process of gradual speciation in which there is a gradual genetic divergence between species populations.
Involves interaction of heritable variation, natural selection, and spatial separation (subdivision). The
classical model of speciation, allopatric speciation, holds that speciation may occur when some geographic
barrier separates populations that show continuous geographic variation. The example of ring species (Insight
Module) shows us that geographic variation within a single species can result in the origin of new species.

First phase: Physical Separation (Subdivision)

Single population (gene pool) is split into two or more spatially separated populations (gene pools) by
geological or ecological process: mountain uplift, rise and fall of land, or vegetational changes. This process
stops gene flow between two groups of populations, allowing genetic divergence to occur as the populations
evolve independently.

Second phase: Genetic divergence and development of reproductive isolating mechanisms (RIMs)

Selective pressures differ in different areas because the ecological conditions differ.
Any mutation or recombination of genes which increases fitness in each environment will be favored,
resulting in genetic/phenotypic divergence. With the passage of time, new alleles would be fixed in each of
the two populations. As the populations become more and more genetically different RIMs may appear.
Isolating mechanisms become more firmly established with continuing evolutionary divergence.

Third phase: Fall of geographic barrier

The geographic barrier "falls" (or one or more populations disperse) and secondary contact occurs after
establishment of some isolating mechanisms, with possible results, depending on 1) duration of separation, 2)
effectiveness of geographic barrier, 3) degree to which RIMs have evolved, and 4) degree to which ecological
divergence (ED) has evolved. When two formerly isolated populations come into secondary contact with each
other, RIMs and ED may be completeor incomplete.

No change (RIMs complete & ED complete; Case 1: BE, p. 80, Fig. 5-4)
Two new species become sympatric without any mutually-induced evolutionary changes. May occur after
long-term separation involving the evolution of complete reproductive isolation with the species having
diverged into different ecological niches.

Experimental studies of Drosophila pseudoobscura by Dodd (1989) have shown that reproductive isolation can
arise in populations that evolve separately.

• experimental populations were reared on two different media for a number of generations:
four on starch-based and four on maltose-based medium
• electrophoretically detectable enzyme differences were show between the two groups, owing
to natural selection associated with the two media
• mating experiments showed that "starch" flies preferred to mate with "maltose" flies, and
"maltose" flies preferred to mate with "starch" flies, i.e. premating/prezygotic isolation
• Rice and Hostert (1993; Evolution 47: 1637-1653) recent review of evolution of prezygotic
isolation in 14 experiments conducted over 40 years: 11 of 14 showed showed significant
change and others showed no significant difference; good experimental evidence of evolution
of reproductive isolation in geographically separated populations

Character displacement (character divergence) (RIMs complete & ED incomplete; Case 2: BE, p. 80, Fig. 5-4)

Differences between species are accentuated in areas of sympatry versus areas of allopatry, which reduces
competition and strengthens reproductive isolation (species recognition), although reproductive isolation
already is complete. Mutually-induced changes in ecological characteristics, which involves behavior, habitat
use, and food selection. It will be strictly ecological at first, leading to different use of the environment; but it
subsequently will be reinforced by selection of morphological differences that facilitate the ecological
divergence (Mayr, 1970). This may occur if geographic separation was relatively short.

Hybrid zone (RIMs incomplete & ED incomplete; Case 3a: BE, p. 80, Fig. 5-4)

A hybrid zone is an area of contact along a common border between two morphologically distinct forms of
organisms. The organisms on either side of the hybrid zone may be races of one species or they may be
different enough to be considered separate species.

When reproductive isolation is incomplete between two species coming into secondary contact with each
other, they may form a hybrid zone along a common border in which hybrid individuals occur. The hybrids
may be both viable and fertile to different degrees. The formation of hybrid zones potentially may allow genes
from one species to flow into another by a process called introgression. One possible outcome is complete
introgression if the hybrids have equal or greater fitness than either parental population. The result is one
species somewhat like the one which existed prior to geographic separation.

Wallace effect (RIMs incomplete & ED incomplete; Case 3b: BE, p. 80, Fig. 5-4)

If hybrids in a hybrid zone show reduced viability or fertility (reduced fitness), natural selection favors further
evolution of reproductive isolation which diminishes hybridization. Natural selection will favor genetic variants
which promote matings between individuals of the same population. Thus, the Wallace effect hypothesizes
that natural selection reinforces reproductive isolating mechanisms.
Reinforcement is the process by which speciation increases reproductive isolation, without regard to the
histories of the populations involved.Secondary reinforcement is the process of reinforcement upon
secondary contact of populations.

The reinforcement hypothesis (in this case secondary reinforcement) maintains that after two allopatric
species subsequently become sympatric slight differences in characteristics associated with mate recognition
are exaggerated by natural selection. The reduced viability and fertility of hybrids owing to genetic divergence
which occurred in allopatry results in selection favoring premating isolation.

Kessler (1966) used Drosophila pseudoobscura and D. persimilis (sibling species) to study artificial selection on
female mate preferences:

• selected females that rejected a heterospecific male to mate with a conspecific male to favor
increased isolation
• selected females that did accepted a heterospecific male to mate with a conspecific male to
favor decreased isolation
• after 18 generations there were significant differences in prezygotic isolation, but there is a
concern about the experiment because it eliminates/ignores the potential effect of gene flow in
natural populations
• Rice and Hostert (1993): eight similar experiments showed similar results

Reinforcement of isolating mechanisms is a controversial hypotheses, and is subject to considerable debate

and testing.

Under the allopatric theory of speciation, reproductive isolation may evolve between two geographically
isolated populations. If it does not the two populations simply merge back into one when they meet again. A
third alternative is that partial reproductive isolation sometimes might evolve during separation and then
undergo reinforcement after secondary contact, which probably operates with the strongest force soon after
the populations meet again. Both scenarios involving reproductive isolation are consistent with the allopatric
theory of speciation. Simply put it holds that geographic separation leads to the fixation of different alleles
through natural selection and to reproductive isolation. The two major alternative modes of speciation --
parapatric and sympatric-- each are heavily dependent on the process of reinforcement.

The Peripheral Isolate Model.-- The process of allopatric speciation we have discussed has been called
the dumb-bell model by Mayr. Under this model, the parental population is divided into roughly two equal
halves. The peripheral isolate model of speciation involves a small population that becomes isolated at the
extreme limit of a species' range. The same sequence of evolutionary divergence and subsequent contact
could occur as was described for the dumb-bell model. The main difference is the relative size of the two
populations.

Mayr and others have argued that speciation via peripheral isolates has occurred more often that
through subdivision. Two arguments in favor of this possibility are: 1) formation of peripheral isolates may be
more likely than physical subdivision of an entire species' range, and 2) distinct forms of a species frequently
have been observed among isolated populations at the edge of the species' range, with individuals in the
center of the range being more similar. The general occurrence of distinct peripheral isolates has not been
demonstrated by systematic study as we only have examples in the current literature (Ridley, 1996). If the
divergence of peripheral isolates from parental populations is shown to be a general pattern, speciation
through peripheral isolates may have been common.

Parapatric Speciation

Species divergence without geographic barriers. Genetically unique individuals exploit contiguous but slightly
different environments (new niches).

Parapatric populations occur in separate geographic areas but share a common border (hybrid zone, primary)
along which hybridization and gene exchange may occur. In the hybrid zone, a stepped cline exists and
reproductive isolation is increased by reinforcement as natural selection operates against the hybrids.

Common in plants, especially Clarkia, due to chromosonal rearrangements. Also, possibly found in some
rodents living below ground and having little mobility, e.g. pocket gophers in rocky mountains (Thomomys
talpoides).

Example:

Clarkia biloba and Clarkia lingulata: Species are separated by extensive chromosomal rearrangements,
including translocations, which form a sterility barrier. Also an example of abrupt speciation (BE: pp. 90-91 &
Table 5-3, p. 91)

Stasipatric Speciation

Another model of parapatric speciation which is very controversial and not widely accepted (Futuyma).

Incipient species arise within range of parent species (sympatrically) by means of chromosomal
rearrangement, and partial reproductive isolation occurs immediately.

The population with the new karyotype spreads from place of origin and establishes parapatric distribution.
Competition between the two forms ensures non-overlapping (parapatric) distribution.

Selection against heterozygotes results in perfection of reproductive isolation through additional RIMs.

Example:
Wingless (flightless), morabine grashoppers of Australia in genus Vandiemenella (BE, p. 86-87). Species with
different chromosomal morphologies (karyotypes) are found in adjacent territories. Different karyotypes are
the result of chromosomal rearrangements.

Sympatric Speciation

Divergence without geographic separation. Involves the rapid establishment of genetic divergence,
reproductive isolation, and new niches in a new species population within the range of a parental population.

This process is difficult to explain in sexually reproducing species. It is very controversial and is not widely
accepted.
This process may, howver, explain the evolution of large genera of plant-feeding
(phytophagous), monophagous (single-host) insects such as the fruit flies inRhagoletis, the apple maggot (BE,
p. 82-85). These cases involve host shifts due to gene mutations resulting in different host selection
(premating mechanisms).

Sympatric speciation does occur in plants

- often through polyploidy (multiplies of chromosome sets)

- example of quantum speciation

Abrupt Speciation (Saltational Speciation, Quantum Speciation)

Saltation = sudden leap across discontinuity.

Instantaneous or sudden speciation through single individual or offspring of single mating is reproductively
isolated from parental stock and is reproductively and ecologically capable of establishing a new species
population.

Two hypothesized mechanisms:

1. Genetic (genic) mutations: not considered possible, only of historical interest

2. Chromosomal changes

Chromosomal changes

Only accepted theory of saltation; supported by empirical studies.

POLYPLOIDY -- only unequivocally established mode of abrupt speciation.

[Note: BE, p. 90, also lists shift from outcrossing to inbreeding in Gilia of California as second
mechanism of quantum evolution plus catastrophic selection via flush-crash cycles and genetic drift.]

Major mode of speciation in plants

Polyploidy: multiplication of chromosome sets (> 2 haploid sets of chomosomes)

Polyploids are reproductively isolated from ancestial species and are new species

AUTOPOLYPLOIDS

More than two haploid sets of single species form zygote

Several examples in BE, p. 93.

ALLOPOLYPLOIDS (AMPHIPLOIDS)

Chromosome sets of two species are combined in hybrid

 Chromosome number is doubled in hybrid yielding a new amphiploid species

Results in hybrid speciation (termed secondary speciation in text)

Most important form of polyploidy

Examples: BE, p. 92-93 (especially Fig. 5-16)

Rare in animals

Parthenogenic lizards and fish arose by hybridization and polyploidy.

Sexually reproducing suckers (catostomids) and salmon too! (tetraploids)

Polyploidy provided genetic material for major adaptive changes.

Most often in plants

Associated with sex-determining mechanisms

Many species have arisen by polyploidy

Estimates of 33%-(36%)-40% of plants originated by such means (intrageneric estimate =

speciation events since genus arose)

If one considers all species in which polyploidy has occurred in the phylogenetic background
(paleopolyploids), the estimate of such species is much greater, approaching 55 percent of
plant species (monocotyledons and dicotyledons).

Example: The tobacco plant, Nicotina tabacum, resulted from doubling of chromosomes of
hybrid between Nicotina otophora andNicotina sylvestris.

EVOLUTIONARY THOUGHT BEFORE DARWIN

CULTURAL TRADITIONS

The intellectuals and naturalists who have considered evolution through the ages have been influenced by the
philosophical movements through history.

Materialism emphasizes the physical world and is the philosophy that matter is the only thing in the universe
which has reality. Matter is, therefore, the basis of all that exists. For example, materialism holds that all
mental processes are caused by physical changes in the body and nervous system; materialists deny the
existence of mind or soul which is distinct from matter.

Classicism stressed order, balance and simplicity in life -- in addition to reason, logic and analysis -- in seeking
that which is universally true, beautiful and good. Classical movements in the west occured first in ancient
Greece (reaching its height in 400's - 300's BC) and continued into the 1600's - 1700's in western Europe.
Romanticism stressed imagination, inspiration and emotion (passion) in seeking the exceptional and
unconventional. The romantic movement occurred in the 1700's - mid 1800's. Romanticism contrasts with
classicism, and often was a revolt against classicism.

SCHOOLS OF THOUGHT

Ideas about evolution existed long before Charles Darwin even appeared on the face of the Earth. Prior to
Darwin's treatise, On The Origin of Species ..., there were several schools of thought regarding evolution. We
will consider the schools and their philosophical views in chronological order to understand the major
philosophical views leading up to Darwin's publication of On The Origin of Species ....

MATERIALIST SCHOOL

The Materialist School began more than 2000 years before Darwin with Thales (640? - 546? BC), who was the
founder of Greek philosophy. It was concerned with material change, and produced first ideas about
processes of evolution. For example,

Empedocles' (495? - 435? BC) ideas presaged the concept of adaptation.

Lucretius (94? - 55? BC) was a Roman poet who formalized materialist views in De Rerum Natura ("On the
nature of things", circa 55 BC): He apparently had a general understanding of the fundamental aspects
of natural selection and adaptation, but he did not understand the mechanism. He also believed inextinction.

CLASSICAL SCHOOL OF THOUGHT

Eclipsed Materialist Tradition 3-4 BC, but demise of Materialist School did not occur till about 1 BC.

Emphasized final, static perfection in organisms.

Complex school of thought comprising many ideas contributed by a number of well known philosophers and
naturalists over about 2,000 years.

Two important ideas: Fixity of Species and Scala Naturae.

Fixity of Species

Fixity of Species -- concept that each species remains unchanged indefinately after its creation. Species were
viewed as discrete, fixed entities which were sharply distinguished from other species and invariable. This
concept was consistent with the views of catastrophists, creationists, and progressionists.

Plato (427?-347 BC): Theory of Forms (Theory of Ideas)

Two worlds: 1) perfect world of Forms and 2) imperfect world. Eidos is the eternal, perfect, heavenly
embodiment of things. Variation in living and non-living things are imperfect manifestations of perfect world
of Forms, and as such are not important.

Aristotle (384-322 BC): Theory of Types, application of Plato's Theory of Forms to biology.

Species reflect existence of unchanging, ideal form, the "universal" or "type", and variation represents an
imperfect manifestation of underlying type. Hence, variation was not all that important--just "noise".

Carl Linnaeus (Swedish botanist, 1707-1778)

His work represents best of Classical Tradition.

Father of Taxonomy, wrote Systema Naturae ("Natural System") and established binomial nomenclature.

Views were decidedly classicist; each species was immutable creation of God, which reflected an unchanging
heavenly type. He believed that an entire species could be represented by one type specimen (holotype). This
view of species is now referred to as the Typological Concept of Species. It stressed
the constancy and discreteness of species, which were considered to be invariable and sharply different.
Systematists of today designate types (e.g., holotypes) when describing species today as a vestige of this view
of species.

These ideas of Plato, Aristotle and Linneaus are referred to as the concept of the fixity of species. Under this
concept, species are viewed as fixed, unchanging entities. Once species originated (were created) they were
believed not to change. These views are entirely at variance with Darwin's views as he consideredindividual
variation to be of great importance and believed that species changed through time, resulting
in transmutation (nowadays speciation).

Scala Naturae ("Scale of Nature")

Another concept of Aristotle.

All creatures arranged on fixed, unchanging linear scale (hierarchy) of nature from lowest to highest forms,
representing degrees of perfection. This idea also is called the chain of being, scale of being, or ladder of
perfection: involved greater degrees of perfection of species with humans on top.

Later in 1600-1700's, Lamarck modified the idea of scala naturae from a fixed scale of perfection into
continuous moving scale of progress, e.g. stair vs. escalator.

This idea too is entirely contrary to Darwin's views as he envisioned that descent with modification would
result in branching evolutionary patterns. Moreover, distinguishing among higher or lower forms of life
or degrees of perfection was not part of Darwin's theory.

DEMISE OF CLASSICAL SCHOOL

A diverse mix of factors eventually led to the demise of the Classical School which dominated thought for a
very long period of time. Environmenatlism wasperhaps the most important factor in bringing the Classical
School to its end.

Environmentalism

Environmental determinism: environmental determination of traits; environmental influences on behavior

(migration).

Although not limited to France, it florished there.

ADAPTATION was a focus of Environmentalist School, especially in 19th century: various mechanisms
proposed.

Climate was considered a major factor of the environment. For example, Buffon noted that animals were
suited to their environments and viewed them to be the products of the climate they experienced.

The eventual failure of the Environmentalist School was its inability to explain the mechanism for the effect
of the environment in determining traits.

Etienne Geoffroy Saint-Hillaire (Fr., 1772-1844)

All adaptations are directly induced by environment and passed on to offspring; focused on influences of
temperature in producing adaptations.

Jean Baptiste de Lamarck (1744-1829)

Philosophie Zoologique (1809): presented theory of evolution.

Believed in scala naturae but turned it into continuous, moving scale of progress toward perfect form versus
the static scale of perfection.

Emphasized continuity of living organisms: species progressed up escalator from spontaneous generation to
pinnacle with man at top through a process of transformation of one species into another.

Believed in evolution (transmutation of species); oddly did not believe in extinction.

Adaptations originated through the use of an organ as a result of an animal's own volition in responding to
adverse, unfavorable environmental conditions. Acquired traits were passed onto offspring if present in each
adult of a mating pair.

Inheritance of acquired characteristics was not new to him.

Charles Lyell (see below) spent great deal of effort refuting Lamarck's ideas (Burkhardt, 1984), and in the
process, formulated the species problem that Darwin solved (Hull, 1984).

NATIONAL SCHOOLS
In the early 1800s, a number of national schools, each with its own traditions, arose and replaced the Classical
School of thought. The separate schools of thought were the result of political turmoil which limited the
exchange of ideas among scientists. There were three main schools of thought:

France -- Environmentalist School: Environmentalism

England -- Natural Theology

Emphasized purposeful design in nature.

Teleology - structure determined by functional results. Explanation of phenomena/processes by purposes

(goals, ends) they serve; purposeful determination by future goals.

"Argument from design" -- design in nature is evidence of benevolence, omnipotence, and existence of God
as Creator of the existing order -- formalized by Thomas Aquinas.

Rev. William Paley (1743-1805; 1802: Natural Theology, or Evidences of the Existence and Attributes of the
Deity): The foremost book of this school, which was widely read at Cambridge including by Darwin. Many
examples Paley provided for his arguments also were used by Darwin because they were cases of remarkable
adaptation.

Germany -- Naturphilosophie

Romantic school: "Bauplan" -- central concept of writers who emphasized the "unity of Creation" with little
emphasis on adaptations of individual species. Bauplan is a common structural plan of life, e.g. skeleton of
vertebrates built on similar plan.

CONTRIBUTIONS OF GEOLOGY

Advances in geology near the end of the 18th century and in the early 1800s made a number of contributions
to our understanding of the Earth's history and to Darwin's ideas.

Geology initially was tied closely to cosmology which addressed the nature and origin of the universe and was
linked to biblical teachings. For example, John Ussher (Irish, 1581-1656) was the first to propose age of Earth
in years by using the Bible to add up the years. He concluded the Earth was created in 4004 BC.

Three major areas of debate in geology were:

1. Reality of Fossils
Medieval scholars considered fossils either to be, 1) products of inorganic processes or 2) false signs of
past life placed in the rocks by Satan to lure people from faith in God or by God to test one's faith.
2. Age of Earth
Until the work of Charles Lyell (see below), the age of the Earth was not considered to be very great.
3. Possibility of Species Extinction
The was a debate as to whether species extinction did or did not occur.
James Hutton (Scottish, 1726-1797) & UNIFORMITARIANISM

FOUNDER OF UNIFORMITARIANISM
First published ideas in Theory of the Earth (1788)
Contrast to views of catastrophism.

CATASTROPHISM

Related to arguments over age of the Earth.

Current rates of geological change rejected as having existed throughout all time, allowing argument that
Earth is young, not old.

Earth's features, including the stratigraphic ones, were postulated to have resulted from extensive, rapid,
violent changes (catastrophies), which were interspersed with long periods of little change.

Catastrophists argued for a young Earth because rates of change had been much greater in earlier periods.

UNIFORMITARIANISM

Counter to "mosaic geology," catastrophism, and progressionism. (Mosaic geology is the account of Creation
in Genesis.)

Rates of geological change are constant through time and extremely slow, requiring incredible time spans to
effect changes. Thus, held that the Earth was old.

Observations of stream erosion convinced Hutton of extremely slow geological processes, requiring a very old
Earth to complete them as processes occured at same rates in the past as in the present day.

Georges Cuvier (Fr., 1769-1832) & CATASTROPHISM

A paleontologist, founder of comparative anatomy, and forceful proponent of catastrophism--despite his

studies of anatomy of fossils which led him to believe in both order and close relationships between living and
dead organisms (unity of nature vs. the pandemonium expected in fossil record associated with
catastrophism).

He did not believe in evolution, but he believed in the reality of fossils and possibility of extinction. Along with
others, he established the reality of fossils. Thus, his own theory of the Earth's history had to account for both
the existence of fossils and the occurrence of extinction.

His theory: The Earth's history was a series of cataclysmic events, with each stratum laid down by a
catastrophic flood or earth quake. In each catastrophe, all life of the period was destroyed and buried in a new
stratum as God destroyed and recreated each of the four major types (phyla) of animals he recognized
(Radiata, Mollusca, Articulata, Vertebrata). God did this because not satisfied with prior creation and made a
better one.
PROGRESSIONISM -- belief in successively better creations. After each cataclysmic episode, each new
creation brought an improvement over the previous one and all such creations were based on a "Divine Plan,"
as manifested in the unity of nature. There was a progression from simple to complex forms of life through
geological time. This idea meshed well with Cuvier's findings and the ideas of catastrophism and creationism.

His theory was contrary to the scala naturae by recognizing four different branches of the animal kingdom
which could not be arranged on a single hierarchy (hierarchies possible within). The rejection of the single
hierarchy led to the demise of the classicist school.

Charles Lyell (Scottish, 1797-1875) & UNIFORMITARIANISM

Principles of Geology (Vol. 1, 1830; Vol. 2, 1832) had a significant influence on Charles Darwin's thinking.
Darwin had copy of Principles of Geology (1830) with him on Beagle (1831).

Responsible for change from CATASTROPHISM to UNIFOMITARIANISM.

Lyell's arguments for UNIFORMITARIANISM:

1. Natural laws do not vary through time.

2. Geological features were created by processes we now see operating, i.e. the same
forces at work today shaped the Earth in the past. Thus, present-day events are a key to
the past and can be used to infer past events.
3. Rates of geological processes did not vary with time (no longer believed). Hence,
cataclysmic explanations were not necessary.

NONPROGRESSIONISM

Lyell's theory that there could be no progress or overall change under conditions of uniformitarianism.
Contrary to Cuvier's theory: there could be no successive series of creations. Contrary to Lamarck's theory:
there could be no transformation of one species into another.

Lyell initially did not believe in evolution. He attacked Lamarck for belief in evolution and also for denial of
extinction. In Volume 2 of Principles of Geology, he used ideas we now know as stabilizing (normalizing)
selection to argue against evolution. He initially criticized Darwin, but later accepted Darwinism in last few
editions of Principles of Geology.

Vestiges of the Natural History of Creation

Robert Chambers (1802-1871) Wrote Vestiges of the Natural History of Creation anonymously in 1844.

The book addressed main scientific issues of the period, including a well-written explication of a scientific
theory of origin and subsequent evolution of life on Earth. It was read carefully by Darwin, who had
formulated his theory of transmutation about 1838.
The book drew strong, vigorous criticism, perhaps decreasing some later anti-Darwinian attacks. The criticisms
also may have caused Darwin to wait more than ten years to publish his theory. They also may have caused
Darwin to present his arguments in moderate, well-reasoned prose.

MAJOR IMPACTS ON DARWIN'S THINKING [chronological order; BE, p.26-32]

1. Lyell: Principles of Geology, volume 1 (1830) -- uniformitarianism, see above

2. Voyage of Beagle
a. St. Jago, Cape Verde Archipelago: possibly convinced Darwin about uniformitarianism.
b. Pampas near Montevideo: fossils.
c. Galapagos Islands: most important area -- animals, especially Darwin's finches which
showed gradation and diversity of structure suggestive of one species radiating into
many.
3. Malthus: An Essay on the Principle of Population

Read by Darwin in October, 1838, about the time he formulated the theory of transmutation.
Before reading Malthus, Darwin believed in the natural theological idea of perfect adaptation.
Darwin shifted to the ideas of relative adaptation and theprinciple of divergence after reading
Malthus. Many biographers believe this change occurred about 1838, but Ospovot (1981) has
argued that Darwin continued to believe in perfect adaptation in the same way as his
contemporaries did until 1854. Ospovot questions the nearly 20-year delay (1838-1854) in this
transition and suggests it was either fear of ostracism or that Darwin was still working on his
ideas. Ideas of perfect adaptation and harmony in nature were linked in the minds of natural
theologists. Relative adaptation meant that natural selection was operating continuously.

SUMMARY: IDEAS IN DARWIN'S TIME

When Darwin wrote On the Origin of Species..., the prevailing, orthodox view of species was that they were
non-changing entities (species fixity). Darwin's claim that species evolved into other species then was
immediately controversial.

Other writers in Darwin's time and before (including his grandfather) had questioned the idea of species fixity.
These ideas primarily addressed thepossibility that species may change into other species, but Darwin's
theory explained the mechanism (how, why) for species change and, in doing so, explained their evolution
into other species.

Jean-Baptiste Lamarck (Philosophie Zoologique) was the most influential of the pre-Darwinian authors to
argue that species change over time.Lamarck's and Darwin's ideas were very different. Lamarck considered
the "transformation" (historians' usage) of species lineages which indefinitely changed from one form into
another and did not become extinct (vs. Darwin). The lineages did not branch, as did Darwin's.

Two mechanisms were used to explain the transformation. The first was that some unknown, internal
force/mechanism would cause an organism to produce offspring which were slightly different from itself.
Accumulation of these changes would then lead to the transformation of a new species.
The second mechanism was the inheritance of acquired characters ("Lamarckian inheritance"), which was not
new to Lamarck(ancient idea extending back to Plato) and erroneously attributed to Lamarck (as the essence
of his theory) by Weismann (Germplasm Theory). During the development of an organism, it acquired
characteristics as a result of exercise. Species could be transformed through the inheritance of individually
acquired characteristics. Volition ("striving", "wishing", "willing") of the organism has been attributed to his
theory. But some authors contend that volition is not a requirement of the theory. Instead, there must be
flexibility in development coupled with the inheritance of acquired characters.

Cuvier was Lamarck's primary antagonist. Cuvier's responses to Lamarck helped to establish in the minds of
biologists the idea that species were fixed, unchanging entities. Cuvier and his associates (school) studied the
anatomy of animals to discover the fundamental, fixed plans by which they were designed. Cuvier also
established that extinction does occur.

Cuvier believed each species had a separate origin and remained an unchanging, fixed form until it became
extinct. This fixity of species was the prevailing view at the time Darwin's work appeared. Catastrophism and
Cuvier's idea of progressionism also were popular views of the time.