Midterm 1 - Lectures Flashcards

(447 cards)

1
Q

Speciation

A

The evolutionary process by which new biological species arise. More specifically, when species reproductively isolate and diverge.

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2
Q

Allopatric speciation

A

Groups from an ancestral population develop into separate species due to a period of geographical separation (lake or canyon).

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3
Q

Sympatric speciation

A

Groups from the same ancestral population develop into a separate species within the same geographic borders.

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4
Q

What did early naturalists believe about speciation?

A
  1. Understood extinction
  2. Thought species creation ended in the Garden of Eden
  3. So, they believed that we were continually losing species without gaining any
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5
Q

How many phases of speciation research?

A

Three

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6
Q

Explain phase one of speciation research.

A

This involved Darwin’s Origin of Species book.

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7
Q

What did Darwin’s Origin of Species book focus on?

A

More on the change within species than the origin of new species. He also recognized that species evolve and divide.

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8
Q

What were the two successors of Darwin?

A

Naturalists and mutationists.

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9
Q

What did naturalists reject from Darwin?

A

They didn’t reject a claim, but thought he focused too much on sympatric speciation since most closely-related species tend to be allopatric.

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10
Q

What did naturalists agree upon Darwin?

A

They agreed with Darwin’s belief that natural selection was the most important force in speciation.

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11
Q

What did mutationists reject from Darwin?

A

They rejected his claim that speciation was gradual and driven by natural selection.

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12
Q

What did mutationists believe?

A

That speciation was separate from natural selection and instead speciation involved nonadaptive and macromutational leaps.

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13
Q

When did phase 2 of speciation research begin? By who?

A

In 1935 by Theodosius Dobzhansky, producing the modern synthesis period.

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14
Q

What did Theodosius Dobzhansky publish?

A

A Critique of the Species Concept in Biology

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15
Q

What was Theodosius Dobzhansky?

A

A naturalist and geneticist.

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16
Q

What did Theodosius Dobzhansky see about alleles?

A

That a continuous evolutionary process – change in allele frequencies – could produced genetically and morphologically discrete groups living in one habitat.

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17
Q

What did Theodosius Dobzhansky believe was missing from Darwin’s theory?

A

He noticed that ecologically distinct forms cannot co-exist without barriers to gene flow, thus stressing the importance of reproductive isolating mechanisms.

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18
Q

What did Theodosius Dobzhansky produce in 1937?

A

The Genetics and the Origin of Species

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19
Q

What was the importance of the Theodosius Dobzhansky’s Genetics and the Origin of Species book?

A

It was the first rigorous genetical studies of reproductive isolation that inspired new research on speciation.

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20
Q

What did Ernst Mayr publish in 1942?

A

Systematics and the Origin of Species

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21
Q

What was included in the Systematics and the Origin of Species book?

A

The biological species concept and the idea that species only arose from allopatric populations.

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22
Q

Biological species concept

A

Organisms belong to the same species if they can interbreed to produce viable, fertile offspring. They are reproductively isolated from other groups.

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23
Q

What did Mayr’s ideas help with?

A

They stimulated research and shaped our current view of speciation.

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24
Q

What did modern synthesis focus on?

A

Reproductive isolating barriers (NOT how the isolation evolves).

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25
What did theoretical and experimental geneticists focus in the modern synthesis era?
Changes within species rather than the origin of new ones.
26
What did evolutionary biologists do in the modern synthesis era?
Focus on using DNA technology to characterize variation within species.
27
What was omitted from the modern synthesis era?
Ethological isolation
28
Ethological isolation
Differences in sexual preferences between species that prevent them from mating.
29
When is the last phase of speciation research?
Began in the early 80s
30
What happened in the last phase of speciation research?
New molecular tools allowed novel and more powerful approaches to old questions.
31
What did the researchers in the last phase realize about the modern synthesis era? Importance?
That ecological approaches to speciation have been neglected, which in return helped in long-term field studies.
32
What did Mary Jane West-Eberhard do in the last phase of research speciation?
Was the first to emphasize the link between sexual selection and speciation. Populations under divergent sexual selection could become behaviourally isolated (no longer recognize the other).
33
Divergent sexual selection
Changes in mating signals and female preferences that leads to another specie emerging, and thus reproductive isolation.
34
What are the important themes in recent research?
1. Continuation of genetic approaches 2. Mathematical theory 3. Emphasis on ecology 4. Molecular analysis 5. Comparative studies (last 4 are novel)
35
What are reproductive isolating barriers?
Those that don't allow organisms to interbreed with one another: - pre-zygotic: prevent mating from occurring like different mating habitats, rituals, or genitals - post-zygotic: reduce the viability (in viability) of the zygote or the fitness of the offspring (sterility)
36
Why was Darwin's ideas controversial?
Went against the idea that all species were created from God's period as he believed that species can constantly change and evolve.
37
Naturalist
Someone who observed organisms in the wild.
38
Are species real (explain through different scientists)?
We act as they are real: - naturalists label specimens - systematists reconstruct relationships among species - population geneticists measure DNA variation within species - ecologists calculate species diversity and document extinct species
39
Why do botanists claim species aren't real?
They claim species are subjective divisions nature for human convenience. This is because mating systems that plants have make it more difficult to consider a species' boundaries
40
Did Darwin believe species were real?
He didn't think they were real and he believed that the classification of organisms was a matter of convenience and not an objective way of summarising diversity in nature.
41
What did Coyne and Orr name as contradictory?
That evolutionists who doubted the validity of species acted as they were real when doing their own research such as using the Linnaean names.
42
What are three methods to determine whether species are real in sexually reproducing groups?
1. Arguments from common sense 2. Agreement between 'indigenous' and 'scientific' species classifications (compare those long time ago that aren't trained in species) 3. Statistical identification of clusters (measure traits of certain animals and see whether the ones you would normally group together as a species, cluster together)
43
What is the first argument that species are real?
They are real because everyone recognises they are real. Dobzansky quoted that formation of discrete groups is universal that it must be regarded as a fundamental characteristic of organic diversity.
44
What is the second argument that species are real?
Clusters are often discrete even to causal observers - there is no continuum between eagles and crows or hummingbirds and parrots (so distinct)
45
What is the third argument that species are real?
The usefulness of field guides is proportional to the discreteness of taxa (can identify the differences between different species).
46
Counterargument for common sense of species being real.
Humans have the tendency to divide a continuous array of organisms into discrete units. ex. dividing the continuous wavelengths of a rainbow (shorter; violet to longer; red) into discrete colours
47
How did the indigenous classification of species compare to scientific ones from Mayr's case?
Indigenous people from Arfak Mountains of New Guinea had 136 names for 137 recognized Linnaean species of birds.
48
How to prove a scientific phenomenon?
If multiple different observers can observe the same phenomenon, can prove the validity of it.
49
How did the indigenous classification of species compare to scientific ones from Diamonds's case?
Found 80% correspondence with Fore people of New Guinea and Linnaean birds. They were even taken to lower elevations and asked to name birds that they had never encountered and 90% of 103 Linnaean species were recognized as distinct. Therefore, the correspondence from two dissimilar classification systems show the objective reality of the species.
50
How did the indigenous classification of species compare to scientific ones in plants?
The Tzeltal people of southern Mexico had 471 indigenous names for plants in their area which were 66% identical to the Linnaean species.
51
Why was there a lower percentage of correspondence of the plants in the indigenous vs. scientific ones?
Geographic variation, so the same species could look quite different in different habitats (giving less overlap). They need to look at the same ones in the same habitat.
52
What is the overall consensus of indigenous vs. scientific classification?
At least 70% correspondence between Indigenous and Linnaean species in various studies. This strongly supports the view that people of different scientific backgrounds recognize similar units of natural diversity.
53
What is not a human classification?
DNA
54
How do genetic basis compare to human classification?
The appearance classification connected to the morphology (measuring plumage, recording songs) line up perfectly to DNA recordings.
55
How do animals show the distinction between species?
In many interactions they can identify the correct species: - male robins court the correct female robin - pollinators recognize specific species of plants - host-specific herbivores and parasites recognize specific species
56
Statistical ID of clusters
Measure a bunch of traits from one species and compare those of another species, do they cluster together based on relatedness.
57
What traits do you use for statistical ID?
Morphological, behavioural, reproductive, and molecular traits.
58
What did Neff and Smith find (statistical ID)?
Using discriminant analysis of morphology in sunfish and shiners showed that sympatric species were well differentiated (clustered together) and the hybrids were intermediates between clusters.
59
How does statistical ID relate to non human classification?
Strong similarity between species classification by morphology and DNA.
60
Do birds and flies just form more discrete clusters than plants?
Mayr studied vascularized plants in Concord Woods Massachusetts and 838 species, 80%, were easily distinguished by morphology (9% of named species did not correspond well).
61
Does the presence of hybrids refute the distinctness of species?
No necessarily: - usually rare and sterile - frequency of hybridizing plants is less than 6%
62
Why do plant taxa have less distinct species boundaries?
Due to variable mating systems.
63
What groups fall into discrete categories in sympatry?
Sexually reproducing ones
64
Are higher taxa real?
Most biologist believe species are real but the higher taxa are not because these groups are less distinct making it more difficult for a normal person to distinguish between the two (are a hummingbird and parrot related).
65
How does the indigenous classification compare to Linnaean taxonomy occcur above species level?
It breaks down
66
What did the Tzeltal people of Mexico show in their classification of trees?
The subdivision of trees into vines, grasses, and broad-leafed herbs highlight evolutionary convergence among distantly related taxa (not close relationships). Some that look similar and not genetically related, so higher order classification is more tricky.
67
What did the Rofaifo people of New Guinea show about higher taxa?
Grouped together orders by similarity not functional or economic considerations. Their five group orders are: - hefa (eels, cassowaries, large marsupials, and rodents) - nema (bats, all birds except cassowaries) - hoiafa (lizards, snakes, fish, molluscs, worms, and centipedes) - hera (frogs other than those of three genera)
68
Examples of asexual reproduction
1. Vegetative reproduction 2. Self-fertilization 3. Production of seeds or eggs without sexual reproduction (apomixis or agamospermy; seeds produced from unfertilized ovules).
69
Does complete uniparental reproduction occur? Results?
It is quite rare and leads to genetic clones.
70
Why is there a continuum of variation between sexual forms in plants?
Trade-offs between forms of reproduction where core diploid species undergo sexual reproduction and these species hybridize to form polyploids. Most often then reproduce agamospermy but sometimes can undergo sexual reproduction. ex. dandelion diploids can form hybrids can form octploids (some cannot reproduce with each other)
71
How many core diploid species does the dandelion genus contain (Taraxacum)?
26
72
How many polyploids and derivatives does the Taraxacum genus contain?
2000 microspecies
73
Are these 2000 microspecies different species?
No, they are not all unqiue and isolated from other species. This is because after his 132 classifications of dandelion in British Isles, botanists could only identity 40% of them.
74
How did the identification of the genus Rubus (berries) compare?
Three botanists had three different answers (24, 205, and 494 species). This meant these classifications weren't universally recognizable .
75
What is the best way to establish whether groups are distinct or not?
To perform cluster analysis on many sympatric individuals.
76
What studies identified "distinctiveness" of asexually reproducing individuals?
Sepp and Paal studied the Lady's mantle, which had more than 1000 microspecies. Examined 43 morphological traits across 23 species in Estonia, and the traits all clustered together. Three were distinct (be cautious of these claims because not much lacking statistical support).
77
Microspecies
A group of organisms that are genetically distinct (not greatly) but morphologically similar to others. Can usually arise through asexual reproduction.
78
What does Darwin mean by mystery of mysteries?
Speciation, which is the act of one species becoming two.
79
How do bacteria (prokaryotes) reproduce?
Through asexual reproduction, transduction, transformation, conjugation, and horizontal uptake (naked DNA in their environment). ex. 18% of genes in E. coli acquired this way
80
How do clusters appear in bacteria?
They are distinct. ex. Barrett and Sneath studied 315 strains of the endoparasitic bacterial genus Neisseria. They analyzed 155 phenotypic traits and found 31 distinct phenotype clusters. Clusters included strains with diverse hosts, and the most difficult to distinguish were the sympatric ones (human nasopharnyx).
81
How are species defined?
Based on Mayr's Biological Species Concept and those based on morphological differences (Typological Species Concepts).
82
How many species are there?
More than 25.
83
Why are there still debates about species and its concepts?
Because not one concept is free from ambiguity when it's applied to nature (rare hybrids, geographic variation, etc.); species evolve from other species, and during this process there will be some unclear cases; and no single species concept can encompass sexual taxa, asexual taxa, and those with mixed models of reproduction.
84
What is the purpose of species concepts?
1. Help us classify species in a systemic manner 2. Correspond to discrete entities we see in nature 3. Help us understand how discrete arise in nature 4. Represent evolutionary history of organisms 5. Apply to the largest possible number of organisms
85
What is one of the most important species problem?
Why do sympatric, sexually reproducing organisms fall into discrete clusters?
86
What was the observation behind the biological species concept?
The observation of discrete sexually reproducing groups in sympatry immediately suggests a species concept based on interbreeding and its absence. Recognized by Dobzhansky.
87
Mayr's Biological Species Concept
Species are groups of interbreeding natural populations that are reproductively isolated from other such groups.
88
Reproductive isolating barriers
Biological features of organisms that impede the exchange of genes with members of other populations.
89
Premating isolating barriers
1. Behavioural isolation 2. Ecological isolation (habitat; temporal; and pollinator) 3. Mechanical isolation 4. Mating system isolation
90
Postmating, prezygotic isolating barriers
1. Copulatory behavioural isolation 2. Gametic isolation (noncompetitive or competitive)
91
Postzygotic isolating barriers
1. Extrinsic (ecological inviability and behavioural sterility) 2. Intrinsic - hybrid inviability - hybrid sterility (physiological and behavioural)
92
Must reproductive isolation be complete?
Limited hybridization may be fine as species status can be a sliding scale: - as gene flow decreases, they become more species-like - when reproductive isolation is complete, good species
93
Why isn't ecological differentiation part of the BSC?
- several instances of one species replacing another close relative in nature (occupy same niche and the role), maintaining species status despite ecological similarity - ecologically differentiated taxa lacking reproductive isolation can fuse in sympatry/hybridize - ecological differentiation is neither necessary nor sufficient for speciation
94
Must reproductive isolation be genetic?
Most isolating barriers are genetic but the geographic barriers that play a role in allopatric speciation are not isolating barriers (initiate speciation process but do not continue it fully). This is because they do not prevent gene flow between sympatric species and do not involve biological differences between taxa (can lead to them involving).
95
How to determine reproductive isolation?
Cannot perform all possible hybridizations of all species, but can infer reproductive isolation from morphological (might not recoginze and mate with one another), chromosomal, or molecular traits.
96
Is speciation reversible?
BSC is concerned with isolating barriers operating at present, but these forms can disappear with a change in environment. ex. human disturbance has resulted in the formation of hybrid swarms in Iris
97
Can speciation be reversible (postzygotic isolation)?
Intrinsic postzygotic isolation is efficient at preventing fusion. Developmental pathways can become less compatible in hybrids (sterility and inviability). As time passes, the probability of reversing all incompatibilities approaches zero as more isolating barriers become apparent.
98
What are the advantages of the BSC?
It helps solve the species problem: 1. Explains the existence of discontinuities among sexually reproducing organisms living in one area 2. Phylogenetic species can be recognized as discrete tips in phylogenies but do not tell us WHY the tips are discrete 3. Suggests a research program to reduce "the origin of species" to "the evolution of isolating barriers"
99
What are the problems with the BSC (allopatric taxa)?
1. Biological species are best diagnosed in sympatry but many taxa are geographically isolated and morphologically differentiated populations (ex. North American elk and European red deer are both Cervus elaphus but differ in size and colour). Testing these predictions would be very difficult 2. In African rift lakes, dozens of cichlids are classified as species based on colour but we do not know if these differences would prevent hybridization in sympatry
100
Are allopatric populations actually separated?
No, as some can exchange migrants and migrants have the ability to interbreed to establish a species status.
101
What is a problems with the BSC (hybridization and introgression)?
Only a problem if one adheres to a strict BSC where no gene flow is permitted.
102
List the attributes of problems of BSC with hybridization and introgression.
1. Much hybridization caused by human disturbance 2. Hybridization may be transient phase during sympatric speciation and reinforcement 3. Some apparent hybrids may be geographic variants or non-genetic variants produced by local conditions
103
Why are Quercus Oaks a problem for BSC?
Is a rampantly hybridized or well differentiated species. Botanists have paid great attention to them.
104
What is the problem with the BSC (species with uniparental taxa)?
BSC cannot deal with species where sexual reproduction is rare or absent.
105
How to address uniparental taxa with the BSC?
Cannot but is the groups form distinct clusters in sympatry, should use species concepts that address the origin and maintenance of these clusters.
106
What is the problem with the BSC (delineating species in a single lineage or fossils or preserved material)?
1. This will be a problem under any species concept as it's somewhat arbitrary but necessary for scientific communication.
107
How to see if changes are abrupt or continuous in allopatric species?
Look at intermediary areas. Two different species in different habitats and no intermediate between them, then physically isolated and different species.
108
When can diagnosing species be less arbitrary?
More predictable when in a single location.
109
How can you judge reproductive isolation with fossils?
Based on discontinuities between phenotypes. Cannot do colours or vocalizing isolating barriers.
110
Problems with the BSC based on evolutionary history.
BSC is severely criticized by systematists (doesn't always apply). Populations of a single species can be less closely related than populations of different species. Evolutionary histories of different genes can differ from history of taxa as a whole.
111
Paraphyly
Group of organisms that share a common ancestor but not all of its descendants
112
What can eliminate ability to detect paraphyly (variation)?
Interbreeding.
113
How to diagnose species as paraphyletic? Is it good?
Using mtDNA. But it can be misleading too.
114
First possibility for why there are species.
Species exist because they are discrete stable states formed by the self-organizing properties of biological matter.
115
What are the issues with the first possibiltiy of why species exist?
It lacks mechanisms to explain such states, does not explain origin of new stable states, and geographic variation in traits casts doubts on the stability of species.
116
Second possibility for why there are species.
Because they fill discrete ecological niches, thus relying on trade-offs.
117
Why does the second possibility of species existing create disruptive selection?
There are no intermediary levels, so hybrids are viewed as unfit and only the extreme phenotypes take suit.
118
Is the second possibility of why species exist independent of reproductive isolation?
No because it relies on isolation from extrinsic barriers.
119
Third possibility for why there are species.
Reproductive isolation is an inevitable result of evolutionary divergence.
120
What does the third possibility of why species exist apply to?
Only those species that are sexually reproducing.
121
What is the third possibility of why species exist related to?
Ecological niches as adaptations can lead to reproductive isolation.
122
What can sexual reproduction lead to?
Anisogamy, which are different sizes of male and female gametes, leading to the possibility of sexual selection. A diverging one can lead to behavioural or gametic isolation.
123
How can bacterial speciation occur?
Through the occupation of new ecological niches through periodic selection or macromutations.
124
To produce clumps, what must clonal reproduction with a tad bit of mutation be accompanied by?
Ecological diversification
125
What does clustering in sexual taxa depend on?
Ecology and reproductive isolation.
126
What does clustering in asexual and uniparental taxa rely on?
Ecological explanation
127
What is the basis of the genetic species concept?
That there is genetic and phenotypic cohesion = within a species, individuals share a significant degree of genetic similarity and exhibit similar observable traits (phenotypes), which helps to maintain a recognizable group identity. This is through gene flow.
128
What is the basis of the ecological and evolutionary species concept?
That there is evolutionary cohesion.
129
What is the basis of the phylogenetic species concept?
Evolutionary history = how organisms are related to each other, including which species they evolved from and which species they are most closely related to.
130
Ecological differentiation
The process by which species adapt to different habitats, which can lead to the emergence of new species.
131
Example of ecological differentiation and the BSC.
Mannequin birds, ecological similar by diet and morphology (similar niches in habitats); however, they occupy different geographic areas (not in the same habitat) leading to different species.
132
What is a key aspect of species?
That they can only be defined relative to other species.
133
What are species a byproduct of?
Evolution within lineages
134
What critics the relative nature of speciation?
BSC
135
Why would we accept the relative nature of speciation?
Every species concept requires comparing different groups of individuals: - morphological distinctness - ecological distinctness - phylogenetic distinctness
136
Pleiotropy (traditional definition)
One gene that affects multiple traits
137
Pleiotropy (speciation definition)
Characters that evolve within one group have side-effect producing isolating barriers with other taxa
138
Direct pleiotropy
Adaptation to a new habitat can automatically lead to spatial isolation.
139
Indirect pleiotropy
Selection for adaptive differences can lead to intrinsic hybrid inviability or sterility.
140
Epistasis (traditional definition)
Effect of one gene depends on genetic background
141
Epistasis (speciation definition)
Genes evolving in one group produce reproductive isolation by interacting with genes evolving in other groups.
142
How can epistasis come about in speciation?
Intrinsic hybrid inviability or sterility OR behavioural isolation due to changes in males in one lineage and changes in females in another.
143
Why would mathematical models and genetic analysis of speciation differ from intraspecific analysis?
Due to complex interactions between genomes of two taxa.
144
What might speciation models show?
Emergent properties.
145
What are the two challenging tasks of understanding the origin of isolating barriers?
1. Which reproductive barriers are involved in the initial reduction in gene flow? 2. Which evolutionary forces produced these barriers?
146
When gene flow is near zero, do isolating barriers stop?
No, they continue to accumulate.
147
Can barriers distort importance? What kind?
Yes they can distort historical importance, especially in sympatric species. ex. monkeyflowers Mimulus lewisii and M. cardinalis overlap in the Sierra Nevada mountains where they have different pollinators in sympatry providing different phenotypic differences
148
When do isolating barriers act?
Sequentially over the life of the organism, beginning with habitat and temporal isolation in adults and ends with hybrid sterility and inviability.
149
What does the addition of isolating barriers do?
Reduces the gene flow that has escaped previous barriers.
150
When do barriers reduce the most gene flow?
The earlier ones even though the later ones may be stronger.
151
What hybrid isolating barrier develops quicker in drosophila and lepidoptera?
Hybrid sterility evolves faster than hybrid inviability.
152
What happens when divergence proceeds between drosophila and lepidoptera?
Both barriers will become complete and hybrids will be inviable.
153
What would seem critically important to speciation of drosophila and lepidoptera?
Hybrid inviability as no hybrids will appear.
154
First importance of isolating barriers
Absolute strength
155
What does absolute strength tell us?
What proportion of gene flow the barrier blocks?
156
What does absolute strength not tell us?
The current relative importance and/or historical importance.
157
What does a large current absolute effect mean?
The isolating barrier played a major role in speciation.
158
What does the strength of a barrier determine?
Its usefulness for genetic studies.
159
Second importance of isolating barriers
Relative strength
160
What is relative strength?
The proportion of gene flow blocked relative to other barriers.
161
How to measure relative strength?
Must measure all possible forms of isolation and the order in which they act.
162
What does relative strength tell us?
The relative importance of barriers that prevent species from fusing.
163
What does relative strength not tell us?
Cannot tell us which barriers caused speciation.
164
What is the most important isolating barrier? Why?
Prezygotic isolation because since barriers act sequentially, it is probably the strongest one to impede gene flow/
165
Example of prezygotic isolation's importance.
Ecological divergence and mate choice assumed to be more important for speciation in Heliconius butterflies, but some species also show evidence of postzygotic isolation.
166
What happens if prezygotic isolation is complete?
There are no hybrids and we know nothing about the strength of postzygotic isolation. ex. two allopatric bird populations diverge in male colour and female preference --> hybrids of intermediate colour cannot find mates --> leads to reinforcement that completes isolation --> with no more hybrids, would be impossible to know that behavioural sterility was critical in speciation
167
What is postzygotic isolation important in?
Via hybrid sterility it is important in restricting gene flow between ancestral species and newly formed polyploid or diploid hybrid descendants.
168
For polyploids to co-exist with ancestors, what must accompany postzygotic isolation?
Ecological prezygotic barriers
169
What can be similar between extrinsic postzygotic isolation and prezygotic isolation?
May involve the same differences in ecology or sexual behaviour that cause prezygotic isolation, such as disruptive selection or sexual selection making hybrids unfit.
170
What is the order of prezygotic barriers?
Habitat isolation --> temporal isolation --> behavioural isolation --> mechanical isolation --> gametic isolation
171
What is the order of postzygotic barriers?
Reduced hybrid viability --> reduce hybrid fertility --> hybrid breakdown
172
What is the importance of postzygotic barriers (hybrids)?
The persistence of distinct taxa in the face of hybridization suggest that hybrids are at a disadvantage. ex. 19% of Lepidopteran species hybridize and the hybrids of mimetic Heliconius butterflies are less attractive to mates and more vulnerable to predators due to being intermediate in colour and behaviour
173
What is the importance of postzygotic barriers (hybrids part 2)?
For reinforcement to increase isolation, must be some postzygotic isolation that makes hybridization disadvantageous.
174
What is the importance of postzygotic barriers (permanence)?
Intrinsic postzygotic isolation is more likely to be permanent. This is because it's difficult to undo hybrid sterility and inviability based on genetic incompatibilities. Selection would have to act on the same exact genes, restoring original alleles.
175
Why is it easy to erase an prezygotic barrier?
Those based on phenotypic divergence can easily be reversed as they involve genes different from the original.
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What is the importance of postzygotic barriers (introgression)?
In separate studies of Drosophila and Helianthus sunflowers, parts of the genome that have not introgressed (mixed) in hybrid zones are those that cause intrinsic hybrid sterility.
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Introgression
The transfer of genetic information from one species to another as a result of hybridization between them and repeated backcrossing (parent and hybrid).
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What do the genes that have not introgressed involved with?
These genes are associated with chromosome rearrangements. Association would not exists if hybrid sterility evolved after speciation.
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What do sex chromosomes that play a large part in hybrid sterility and inviability show? What does this means?
Reduced movement through hybrid zones which provides evidence that postzygotic isolation operated in the past and continues to operate prevent introgression.
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What is the importance of postzygotic barriers (asymmetry)?
Asymmetry between how pre and postzygotic isolation behave beyond F1.
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When do prezygotic and extrinsic postzygotic barriers become weaker?
In F2 generation and backcrosses.
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Why do prezygotic and extrinsic postzygotic barriers become weaker in F2 gen and backcrosses?
Because many hybrids behave like pure species.
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What do genes causing intrinsic postzgotic isolation do in F2 and backcross hybrids?
They increase the sterility and inviability of these hybrids.
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How to tell which barriers originally caused speciation?
Could study geographic populations of a single species to determine which barriers exist in the initial or early species.
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What is a problem with trying to find barriers that caused speciation?
Most populations never become species and even if a single barriers is first to arrive, it's impossible to determine its later influence.
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How to look correctly as barriers causing speciation?
Comparative analysis of multiple taxa at different stages of evolution is more promising.
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List comparative analysis of multiple taxa at different stages of evolution.
1. Sympatric species with few isolating barriers 2. Hybridization following ecological disturbance 3. Isolating barriers associated with chromosome rearrangements 4. Comparative studies of evolution of reproductive isolation 5. Comparative studies of traits promoting speciation
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What is a major barrier in polyploidy plants?
Postzygotic isolation via chromosomally-based hybrid sterility.
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When can postzygotic isolation of polyploidy plants exist?
Only if there is sufficient ecological differences to co-exist with diploid ancestors.
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What are epilachna ladybird beetles isolated by?
Preference for and ability to survive on different hosts.
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What are echinometra sea urchins isolated by?
Gametic isolation and can co-exist because of it.
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When can you infer a barrier's importance in speciation?
If an ecological disturbance causes previously isolated species to fuse or form hybrid swarms. ex. in lake victoria, there is a positive correlation between number of cichlid species and water clarity (male colour only visible in clear water).
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What do chromosomal rearrangements cause?
If the genes causing reproductive isolation reside within chromosome regions that are rearranged, association will persist when species come into secondary contact.
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What does heterozygosity for rearrangements prevent?
Recombination between genes within rearrangements.
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What will reduce recombination hinder?
The elimination of genes by natural selection against hybrids leading to delayed fusion of species and allowing other barriers to accumulate.
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What do comparative analyses show?
The strength of barriers and the time of divergence between taxa.
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Give a comparative analysis example.
Work by Wilson showed that mammals could produce viable hybrids after 2-3 millions years of divergence, frogs after 21 million years, and birds after 22 million years. NOTE: 22 million years would be equivalent of humans and gibbons producing hybrids.
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What did Coyne and Orr study of Drosophilia species show?
That prezygotic isolation appears to evolve faster (develops at 0.75 whereas for the postzygotic one is takes over 1.0). Disparity is due to biogeography and sympatric taxa diverged faster (more pressure).
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What is the problem with the evolution of isolation studies?
There are only a few others; missing barriers in every group; and most measure the barriers in ONLY Drosophilia, darters, and plants.
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What barrier evolves slowly?
Intrinsic hybrid sterility and inviability as there is no recently diverged taxa with sterile or inviable hybrids.
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What postzygotic barrier evolves faster?
Hybrid sterility evolves faster than hybrid inviability.
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What does intrinsic postzygotic isolation evolve?
At different rates in different groups.
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What traits promote speciation?
Animal pollination of plants and those associated with sexual selection in birds and insects.
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Behavourial isolation
Phenotypic differences that stop mating OR they cannot recoginze each other. ex. plummage colour on birds
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Temporal isolation
Mating at different times of the day.
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Pollinator isolation
Plant species have different pollinators that visit them. ex. hummingbird or bee
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Mechanical isolation
Lock and key mechanism - reproductive parts do not fit.
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Mating system isolation
Plant species have different reproductive strategies. ex. agametic or ploidy levels
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Noncompetitive gametic isolation
Heterospecific and conspecific not interacting at the same time.
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Ecological inviability
Hybrid intermediate in morphology so ill-suited to different habitats, might not survive.
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Behavioural sterility
Survived to age of reproduction, but nobody wants to mate with it.
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Hybrid inviability
Cannot survive to maturity due to a physiological defect.
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Hybrid sterility
Do survive to reproduce, but cannot properly reproduce.
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Explain isolating barriers with Drosophilia melangoaster and D. simulans.
200 genes involved in hybrid inviability with each one causes nearly complete hybrid inviability itself. A lot of genes important in reproductive isolation, but only need a few. This makes it difficult to isolate which one caused it first, especially as they evolve and separate.
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Explain isolating barriers using monkeyflowers.
Habitat isolation accounts for 59% of reproductive isolation and pollinator isolation accounts for 98%. Hybrid inviability results in 12% of reduction in gene flow. With all these different barriers, unsure which one evolved first. Any combination of three barriers reduces gene flow to nearly zero (don't need them all, so it doesn't mean they are all important in the speciation process).
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Reinforcement
Natural selection acting on species to provide reproductive isolation.
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Threespine sticklebacks and the importance of postzygotic barriers.
Closely related species that lives in the same lake but segregated by body size and feeding regimes. It would change if the lake becomes uniform, barriers ceast to be important. Postzygotic barriers more important as they involve deep rooted genetic changes.
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What did Coyne and Orr do?
Plotted pre/post - zygotic isolation with genetic distance (how long they had been isolated).
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How did pre/post zygotic isolation compare in other species for the Coyne and Orr test?
Birds very distantly related could still mate with each other (0 postzygotic isolation). This just means that the strength of barriers can differ in groups.
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Allopatric speciaton
Speciation with complete geographic isolation.
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Parapatric speciation
With limited genetic exchange.
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Sympatric speciation
With free gene flow.
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What did type of speciation did Darwin believe?
Sympatric
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What type of speciation did Ernst believe?
Allopatric
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What are the two forms of allopatric speciation?
1. Vicariant speciation 2. Peripatric speciation
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Vicariant speciation
Geographic range of a species splits into two or more large populations.
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Peripatric speciation
Isolated habitat colonized by a few individuals or a small population becomes geographically isolated.
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What is the geographic separation of vicariant speciation?
Due to climate or geology: - formation of land bridge - glaciation - elevation of mountains - continental drift - climatic change - extinction of intermediate populations
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Does speciation occur quicker in vicariant?
Yes, because gene flow is uninterrupted.
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Why is divergence inevitable in vicariant speciation?
They are adapting to new habitats where they can develop new mutations through genetic drift. All this coupled with no gene flow allows isolating barriers to develop.
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What has confirmed how allopatric speciation can occur?
Mathematical models through drift alone, drift and selection, and sexual selection.
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What have experiments showed for allopatric speciation?
Evidence as few as 18 generations for flies showed prezygotic isolation were a byproduct for selection. Postzygotic isolation were low probably need longer generations.
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First line of evidence for vicariant speciation (strongest).
Geographic concordance of species borders with geographic or climatic barriers. ex. Isthmus of Panama
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What happened before the isthmus was formed?
North America and South America were separated before and this allowed species to easily move from one side to another.
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What have the snapping shrimp genus Alpheus showed (Isthmus)?
13 pairs of sister species in Pacific and Atlantic. One species in the past but when the land barrier came in, they separated over time. Only 1% of matings could yield fertile clutches in lab.
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How do angiosperms in eastern Asia and eastern North America show allopatric speciation?
Once connected continents, but when separated a long time ago, the species diverged more recently based on climate change. Cooler and drier climates replaced the forest habitat with grassland, that's why more restricted to eastern areas. Each of genera have one sister species in north America and one in Asia.
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Second line of evidence for vicariant speciation (second strongest).
Allopatry of young sister species (each other's closest relative). Sister species within a group always occur in allopatry even if no geographic barrier.
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Sea urchins sister species example.
Sister species are always allopatric whereas older ones can overlap.
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Third line of evidence for vicariant speciation (middle).
Geographic coincidence of species borders or hybrid zones among different taxa. Can infer speciation in allopatry where a present distribution reflects secondary contact after disapperance.
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What must you know/not know for the third line of evidence for vicariant speciation?
Must know that a barrier used to exist. Must not be an ecotone barrier where species ranges meet (transition between two biomes).
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Example of third line of evidence for vicariant speciation.
In S. America, Pleistocene advance of glaciers reduced precipitation and temperature and produced refugia, small patches of forest. This produced multiple speciation events. Look at Charis butterflies; sister species viewed populations are completely separated (allopatric) or parapatric. Seen seven areas of endemism identical to those derived from phylogenies of reptiles, birds, rodents, primates, etc.
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Example suture zones in N. America.
6 major zones where they contain many hybridizing plants, insects, mammals, etc. These zones correspond to the sites of former geographic barriers (secondary contact between allopatric ones). ex. red shafted flickers found on west and east coasts OR bullock's oriole and baltimore oriole's
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Fourth line of evidence for vicariant speciation (third weakest).
Absence of sister species in areas where geographic isolation was unlikely. Need barriers to produce speciation. ex. Coyne and Price surveyed 51 remote oceanic islands never connected to mainland; found no sister species of bird; only found when species could move to different island (allopatric speciation)
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Fifth line of evidence for vicariant speciation (second weakest).
Concordance between present or past geographic barriers and genetic discontinuities between species (where there was once a barrier, can show genetic differences) - abrupt changes in allele frequencies within species - changes occur in similar locations across species
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What is the problem with the fifth line of evidence for vicariant speciation?
Patterns show that geographic isolation promotes genetic diveregence, not speciation. This is because you are still looking at individuals part of the same species.
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Example of the fifth line of evidence for vicariant speciation.
Glaciation in the past had no pennisula in Flordia, so animals could move freely. Introduction of Florida has created a barrier (bigger during past glaciations), more water after retreated, created a bit more movement. Allele frequencies change from one side of pennisula to another: horseshoe crab, ribbed mussell, etc.
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Sixth line of evidence for vicariant speciation (weakest).
Increase in reproductive isolation with geographic distance between populations.
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Clinal isolation
Remote populations may exchange genes only rarely allowing for fairly independent evolution.
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What is the problem with the sixth line of evidence for vicariant speciation?
If ecology changes geographically, it could be due to local adaptation rather than geographic isolation itself. Therefore, it's not fully due to allopatric speciation
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What are examples of clinal isolation in nature?
1. Strepthanthus glandulosus plants 2. Desmognathus ochrophaeus salamander 3. Tigriopus californicus copepod 4. Timema cristinae stick insects - their genetic distance increases as you move farther distances between their species
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Ring species
Encircles a geographic barrier like a mountain or plateau where ancestral populations gradually expand their range around the barrier. Exchange gene flow freely expect where they meet at the other end of the barrier (genetically/reproductively isolated for so long).
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What do ring species show?
The attentuation (reduction) of gene flow with distance NOT allopatric speciation.
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Why is ring species better evidence than clinal isolation for allopatric speciation?
Clinal isolation can be caused by environmental differences that increase with distance between populations. Cannot make the same argument with ring species since the most reproductively isolated populations occur in the same habitat. Not habitat driving it, genes are.
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What are the criteria of ring species?
1. Ring was founded by one population rather than two or more genetically distinct populations 2. At least one of the terminal populations must represent the most recent range expansion 3. There must be no present or past geographic barriers around the ring to interrupt gene flow 4. Gene flow is extensive between adjacent populations but decreases with distance around the ring
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Give an example of ring species.
Ensatina eschscholtzii in Western USA. Ancestral population in N. Calfronia and moved south through different areas. Genetic analyses suggest that they form a superspecies that diverged in allopatry.
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What is a better example of ring species?
Greenish warbler in Asia, circular distribution around Tibetan plateau. Played back songs of males in different areas to see if the males recognised the species. SHOWED: - Male song recognition decreases with geographic distance - Males do not recognize each other’s songs in terminal populations
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What kind of selection did the warblers show?
Parallel sexual selection across either side of the plateau for longer male songs: - one terminal = increased song length by increasing syllable length - other term = increased song length by repeating shorter syllables multiple times
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What kind of species is the warbler?
Allopatric species because there was a deep genetic split in the ancestral populations. Still shows that reproduction isolation was enhanced by reduced gene flow around the ring.
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What are models of peripatric speciation?
Verbal ones.
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Why is peripatric speciation separate from normal allopatric speciation?
Colonization events have special features that may lead to much rapid speciation due to adaptation to new habitats. Smaller, different genetic makeup (inbreeding maybe) in a new environment. Genetic drift can lead to fixation of novel alleles due to a restricted genetic sample of original population.
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What are studies of peripatric speciation?
Found that one in flies that those in isolation could be achieved after a series of founder events but rarely for a single one. Small sample sizes in other studies count too.
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First evidence of peripatric speciation in nature.
Peripatric speciation on single oceanic islands. Presence of endemic species on oceanic islands whose closest relatives inhabit nearby continents is strong evidence.
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What is the issue with the first evidence of peripatric speciation in nature?
Do not know if they are reproductively isolated.
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When is the evidence stronger for the first evidence of peripatric speciation in nature?
If the island species is the sole representative of genus suggests that being isolated on that island causes a unique speciation event (not allopatric).
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Example of the first evidence of peripatric speciation in nature.
Cocos Island finch is the only Darwin's finch outside the Galapagos and the only member of its genus.
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Second evidence of peripatric speciation in nature.
Peripatric speciation on archipelagos (group of islands).
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Example of second evidence of peripatric speciation in nature.
Hawaiian archipelago has over 400 species of Drosophila. All descended from a single ancestor with more than 50% of species originated peripatrically with higher % for newer islands. Similar for cyanea plants.
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How do island number and number of species related?
As more islands forming, formation of new species (more possibility for colonization events).
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Third evidence of peripatric speciation in nature.
Peripatric speciation in peripheral isolates (populations at an outside edge of the main species distribution).
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What is the difficulty with the third evidence of peripatric speciation in nature?
1. Harder to detect on continents than islands 2. Sister species with abutting (touching) distributions but unequal range sizes can also result from vicariant speciation followed by reduction in range size
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Example of the third evidence of peripatric speciation in nature.
Clarkia biloba and C. lingulata are separated by 100m. Colonized by a population of C. biloba that speciated rapidly due to limited seed dispersal and some self-compatibility.
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Example of the third evidence of peripatric speciation in nature.
Changes in chirality (coiling) in land snails. Right and left coiling morphs are behaviourally isolated because they cannot copulate.
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What did a recent study show for the third evidence of peripatric speciation?
Comparative study showed that recently-formed allopatric taxa had very asymmetric range sizes, consistent with peripatric speciation.
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Parapatric speciation
Speciation between two populations with limited gene exchange.
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What are the models of parapatric speciation?
Clinal speciation and stepping-stone.
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Clinal speciation
A single species continuously distributed across a variable environment. Microadaptations to changes in environment that contribute to speciation (continuous along habitat).
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Stepping-stone
Discrete populations having limited gene exchange.
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What models exist for parapatric speciation?
Verbal and mathematical models.
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What does the success of parapatric models depend on?
Assumptions: - rates of gene flow - population sizes - genetic basis of reproductive isolation ex. small population, less gene flow, and strong genetic basis of reproductive isolation = parapatric speciation possible
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What models of parapatric speciation are more successful?
Stepping-stones one because they are more isolated.
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Example of parapatric speciation.
Experiment of houseflies Musca domestica. Two lines selected for different traits. Found migration between lines at 30% per generation and significant assortative mating (mating within their own line as opposed to the other one) after 38 generations. Can still have speciation with some limited genetic information exchange.
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First evidence of parapatric speciation in nature.
A pair of closely realted species with abutting (close) distributions.
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What is the problem with the first evidence of parapatric speciation in nature?
Weak evidence as you can't distinguish between allopatric speciation.
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What can provide some solution to the problem with the first evidence of parapatric speciation in nature?
Hybrid zones if you can exclude the possibility of previous allopatry.
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Second evidence of parapatric speciation in nature.
Multiple pairs of related species with abutting distributions, like hybrid zones at ecotones. Better because you have multiple examples of it occuring.
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Third evidence of parapatric speciation in nature.
Morphological or genetic incontinuities (changes) at ecotones, especially at hybrid zones. ex. little greenbul in Africa or lizard Anolis rouqet on Martinique
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What is the problem with the third evidence of parapatric speciation in nature?
Shows a necessary step in the process as you're getting morphological change as the environment does change. Don't know if you continue over time if things will continue to diverge or need a barrier for speciation.
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Fourth evidence of parapatric speciation in nature.
Observations of all stages of parapatric speciation in nature
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What is the fourth evidence of parapatric speciation in nature considered?
Considered this evidence of parapatric speciation (see the whole range from diverged to not diverged) but could still be caused by allopatric speciation.
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Example of the fourth evidence of parapatric speciation in nature.
Jiggins and Mallet (2000) surveyed parapatric populations in several species and found every level of reproductive isolation: - from hybrid zones with random mating (complete gene flow) to boundaries maintained by strict reproductive isolation
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Fifth evidence of parapatric speciation in nature.
Biogeographic and phylogenetic evidence of parapatric speciation in a small clade. Closely related “young” species with parapatric distributions.
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Sixth evidence of parapatric speciation in nature.
Historical observations of speciation in parapatry.
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What is the problem with the fifth evidence of parapatric speciation in nature?
Speciation in allopatry and range expansion (ranges separate, but got larger and touched each other) could cause this pattern AND hybridization (secondary contact if previously isolated) could also cause gene mixing making species appear “younger” (similar so diverged more recently than they are) than they are.
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Example of the sixth evidence of parapatric speciation in nature.
Evolution of heavy metal tolerance in plants growing near mines (more concentrated metals closer to mine, and plants developing adaptations). Sharp ecotone between contaminated and uncontaminated. Partial temporal isolation of flowering times provides some reproductive isolation (mate with tolerant ones) but still a lot of gene flow (NOT true species).
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What is difficult about parapatric speciation?
Hard to determine if truely right because even though they appear to be close environment and habitats, it's unsure whether or not they were separated once ago by barriers or range expansion.
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Sympatric speciation
The origin of an isolating mechanisms among the members of an interbreeding population (all occur within the same range).
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Where must sympatric speciation occur?
Within the average dispersal distance or "cruising range" of a single individual.
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What was Darwin's view on sympatric speciation?
He used a morphological species concept, but didn't understand reproductive isolation yet. He just believed that species would arise in sympatry to fill empty niches.
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What was Mayr's contribution to sympatric speciation?
Realized it was very unlikely to occur based on genetics.
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What is the first problem of sympatric speciation?
Antagonism between selection and recombination: - As selection tries to split apart a population, interbreeding continually breaks up gene complexes that produce reproductive isolation Ex. could break up genes for habitat fitness and habitat preference/NOT broken up could enhance speciation in sympatry
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What is the second problem of sympatric speciation?
Coexistence: - Populations must develop sufficient ecological differences to coexist - Implies that selection should in part be driven by selection for ecological divergence NOTE: sexual selection doesn't cut it because even if you look different, you will be competing for the same food and resources
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What are the two models of sympatric speciation?
1. Disruptive sexual selection 2. Disruptive natural selection
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What does disruptive selection mean?
Distribution of varying extreme individuals. Becomes concentrated on end points (biomodal). The intermediates will decrease.
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In adaptive radiations, what do species differ in?
More strongly in sexual traits than ecological adaptations. ex. african lake cichlids
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Adaptive radiation
The evolution of a single ancestral species into multiple descendant species that differ in their habitat, form, or behaviour. This process occurs when a population encounters new and diverse environments, leading to the development of different ecological niches.
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What do models for disruptive sexual selection not show?
No mechanism for the coexistence of new species in the same habitat.
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Are models for disruptive sexual selection good?
Some success, but usually show unrealistic and restrictive assumptions.
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What have recent models of disruptive sexual selection added?
Ecological divergence.
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What are the two types of natural selection models?
1. Discrete-habitat models 2. Continuous-resource models
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Discrete-habitat model
Sympatric species that seek out different niches, are most fit in their own niches, and preferentially mate with members of their own niches. ex. insects that feed and mate on a specific host plant
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What traits does the discrete-habitat model focus on?
1. Niche preference 2. Niche adaptation (more successful in preferred habitat) 3. Assortative mating (mating with certain members)
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Are discrete-habitat models good?
Not appropriate for many species: - sympatric species of African cichlids fish do not show dramatic differences in habitat (only look different)
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Continuous resource models
Based on co-evolution of ecological traits and assortative mating. It splits populations into two sympatric groups, each using a different part of the resource distribution.
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Explain the continuous resource model from Dieckmann and Doebeli.
- Important resource (e.g. seed size) unimodally distributed – Ecologically-relevant trait (e.g. beak size) unimodally distributed – As population grows, frequency- and density-dependent selection favours individuals with more extreme values (larger and smaller beaks); because of competition for average size seeds/beaks – Splitting occurs if individuals with extreme traits prefer individuals with similar traits
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What are the first problems with the Dieckmann and Doebeli model?
- works best when individuals have innate preference for mating with other individuals having similar traits (large beak with large beak) - if mate preferences are not very strong, speciation takes so long that initial conditions probably change before speciation occurs (1 mill generations)
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What did the medium ground finch study show?
- change in mean beak size - when there was a severe drought, beak size increased afterwards to crack large seeds - new species arrived in a decade - over time, beak size decreases due to competition (average) from a larger competitor - new drought, decreased beak size even more - a short scale for variation makes sympatric speciation in the DD model seem less likely (no preferences for mates)
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Did the medium ground finches show mating preferences?
Yes, different beak sizes create different songs that attract mates.
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What are the second problems with the Dieckmann and Doebeli model?
- As the ends of the resource distribution become occupied, less competition for central resources, which should favor intermediate individuals – Could result in a continuum of interbreeding forms rather than discrete groups
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What should be the null hypothesis for speciation?
Allopatric speciation
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Should you rely on models for sympatric speciation?
No, better to judge the likelihood of sympatric speciation based on laboratory experiments and evidence from nature.
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What have some experiments of sympatric speciation showed?
- demonstrated reproductive isolation between selected lines that interbreed
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What is a problem with some of the experiments of sympatric speciation?
They are "kill the hybrids" experiment where hybrids are removed in each generation. This leads to successful isolation, but it is not realistic.
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What did the house fly geotaxis study do (sympatric speciation)?
- applied strong disruptive selection to flies moving up or down from a central chamber - reared apart but allowed to interbreed
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What did the house fly geotaxis study find (sympatric speciation)?
After many generations, flies diverged in geotaxis and mating tests showed strong sexual isolation.
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What did the bristle number of drosophila study do (sympatric speciation)?
12 generations of disruptive selection for bristle number.
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What did the bristle number of drosophila study find (sympatric speciation)?
Flies showed complete assortative mating by bristle number (preferred individuals with same number of bristles).
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What did the bristle number of drosophila future studies find (sympatric speciation)?
Results couldn't be replicated in 19 future studies, most found moderate assortative mating.
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What was the most successful drosophila study? Findings?
By Rice and Salt and they found that individuals mate only with others who choose the same habitat -- "no gene" assortative mating maze.
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What were the three environmental factors that varied in the fly study?
1. Phototaxis (light or dark) 2. Geotaxis (up or down) 3. Chemotaxis (ethatnol vs. acetylaldehyde) Vials with each.
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How did the fly study stimulate selection for extreme habitat preference?
By only choosing flies that choose the extreme habitats to be allowed to breed: 1. Up/dark/acetylaldehyde 2. Down/light/ethanol
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What were the additional selection in the fly study?
Only choose the flies that hatched early from habitat A (Up/dark/acetylaldehyde) and the flies that hatched late from habitat B (Down/light/ethanol).
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How to tell which habitat the fly's parents had chosen?
Used mutations and chemical markers from a fly's eye colour.
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How did habitat isolation look in the fly study?
Complete habitat isolation in 30 generations which meant all flies who had chosen each extreme environment were offspring of parents who had chosen that environment.
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What did the fly study not show?
No evolution of complete habitat preference.
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How did the fly study not show complete habitat preference?
Offspring flies from extreme habitats continued to choose intermediate habitats but were considered lethal and discarded. When the flies were all put in a common environment they did not assortatively mate.
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What was the overall conclusion of the Rice and Salt study?
That habitat isolation can evolve if selection is strong and assortative mating is a byproduct of habitat choice. Strong disruptive selection appears necessary but not sufficient for sympatric speciation by itself.
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Why is allopatric speciation the null hypothesis?
1. Because it's easier than sympatric speciation. Selection or drift acting on geographically isolated populations will eventually produce isolating barriers. 2. There is also strong and persuasive evidence for allopatric speciation. 3. Many opportunities for geographic isolation; over the past 2 million years, there has been 20 major glacial advances and more frequent cycles of temperature changes
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What four criteria must sympatric speciation meet?
1. Species must be largely or completely sympatric 2. Must have reproductive isolation, preferably of genetic origin 3. Must be sister groups (closest relative) 4. Biogeography and evolutionary history must make allopatry very unlikely
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Monophyletic group
All on a common branch, thus diverging/evolving from a common ancestor.
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Microallopatry
When looking at a distribution, they overlap; however, visiting their locations in person, will demonstrate a small geographic isolation.
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How may species on islands show sympatric speciation?
If a small monophyletic group was confined to a small isolated habitat, it would be convincing.
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Example of species on islands showing sympatric speciation. Problem?
Endemic Coleptera and Orthoptera on St. Helena once thought to have speciated in sympatry, but since the species are all flightless, this means they could have speciated microallopatrically.
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What do flightless weevils on the island of Rapa show?
Could have speciated microallopatrically OR on small islets that were connected to mainland during Pleistocene.
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What do lepidoptera on island of Rapa show?
Suggestion of allopatry because of negative correlation between endemism and mobility (less endemic, more mobile they are). The group wasn't endemic, so multiple colonizations could have occured.
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Why do endemic species fail to provide evidence of sympatric speciation on islands?
No sister species on isolated islands, so they could not separate or diverge in sympatry.
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Why are fishes in postglacial lakes considered sympatric speciation?
Some lakes contain closely related fish differing in morphology, behaviour, habitat, or life history. ex. threespine sticklebacks in same lake but have different vegetation, morphology, and assortative mating
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When did endemic fish evolve?
Most likely 15,000 years ago when lakes in N. America ad Eurasia were formed from the last glaciation.
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Problems of fishes in postglacial lakes for sympatric speciation.
1. Lakes connected to rivers and the sea, which could allow from multiple invasions from allopatric taxa 2. Gene exchange after secondary contact (hybridization) can yield misleading conclusion that the fish are sister taxa; hybrids have intermediate genes (need to use nuclear DNA instead of mtDNA)
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What is strong evidence for sympatric speciation in fished of postglacial lakes?
Arctic charr in Lake Galtabol, Iceland - limnetic and benthic morphs are more closely realted to each other than taxa in four nearby lakes - genetic similarity does not reflect current hybridization: share no alleles at one of six nuclear loci and strongly differentiated at other loci
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What are the two types of lakes in African for cichlid fish?
Rift lake = victoria, malawi, and tanganyika Tiny crater lakes = barombi mbo and bermin (much smaller and isolated, so fewer species)
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How are the speciation rates of barombi mbo and bermin?
Quite high, some higher than the larger rift lakes.
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Arguments for sympatric speciation in African lakes in cichlid fish.
1. Hard to envision geographic barriers allowing formation of hundreds of species in only 2 million years 2. Speciation evidence has also occurred in limnetic and deep water forms less likely to encounter geographic barriers (not in open water) 3. Movement between habitats may prevent allopatric speciation (no walls separating habitats) 4. Closely related species differ more in colour than morphology and habitat use 5. Some pelagic (open water) species show no genetic structuring across Lake Malawi; fish are so mobile that allopatry not possible (don't cluster together genetically; huge intermixing population) 6. Monophyletic groups of pelagic species show that speciation occurred in pelagic ancestors
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Arguments for allopatric speciation in African lakes in cichlid fish.
1. Levels of all three major lakes have risen and fallen repeatedly, creating isolated small lakes 2. Many species have highly localized ranges and shorelines have diverse habitats (rocky, sandy, swamps, bays, rivers) 3. Habitat specificity can reduce gene flow 4. Genetic patterns sometimes mirror fragmentation of lakes (cluster genetically to past areas of lakes) 5. Littoral fish have limited migration but occasionally colonize new habitats, promoting allopatric or parapatric speciation 6. Some pelagic species return to littoral zone for spawning which could cause allopatric speciation by habitat segregation 7. Some allopatric populations evolved behavioural isolation during periods when there were barriers to dispersal
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Where is sympatric speciation more convincing in African lakes (cichlid fish)?
More convincing in the small crater lakes because: - Fish differ more in feeding habits and morphology relating to diet – Little sexual dichromatism – Sister species and no evidence of hybridization – Size and formation of lakes suggests no allopatry (not connected to other lakes in the past; formed in tops of old volcanoes) – Meet the four criteria for sympatric species
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How can species be considered islands?
Host-specific parasites occupying different host species could be considered islands.
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What would be stronger evidence for sympatric speciation of host-specific parasites?
Parasites that occupy different parts of the same host species, so they exist within the same geographic range. ex. lice in birds occupying different feathers or even different parts of the same feather
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Are head lice and body lice in humans different species?
No since genetic studies show similarity.
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Why did it look like fig wasps have sympatric speciation?
Different fig wasps had different ovipositer lengths depending on the type of figs they were parasites too. Also, there was temporal segregation (different fig wasps breed at different times of the year).
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Why were fig wasps then thought to be allopatrically speciated?
Since the fig wasps and different adaptations happened on an island chain (Melanesian).
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Host races
Involve two different races of species living on different host plants in the same area (not true species, evidence that speciation is occuring).
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What is an example of host races?
Apple maggot fly that used both apples (introduced by colonizers) and hawthorns (original host; native) as hosts in the US.
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Is the apple maggot and hawthorns sympatric speciation?
No, recent genetic evidence suggests that the apple race might have descended from an allopatric hawthorn race rather than the original one in the area.
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Host-specific species
Already diverged. They are species that live on different hosts and show little gene flow. ex. treehoppers
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Why were treehoppers thought to be sympatric?
Each of six species inhabits a different species of tree and they are reproductively isolated.
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Are lacewings host-specific species that are sympatric?
No, genetics show that they are not sister species. Referring to the ones that differ in colour and habitat.
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Allochronic (temporal) isolation
Different times of breeding or flowering.
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Are gryllus field crickets sympatric?
Even though the two species breed at different times, they are not sister species.
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What species is an example of allochronic isolation in sympatry?
Periodic cicadas that breed in 13 year vs 17 year cycles shows influence of rapid changes in life cycle of reproductive isolation. NOTE: cannot exclude biogeography
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What do comparative studies do?
Calculate degree of range overlap across groups of closely-related species.
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What do comparative studies of allopatric speciation show?
Changes in range over time should lead to slightly increased overlap.
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What do comparative studies of sympatric speciation show?
Range overlap should decrease over time (since speciation occurred during complete range overlap).
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What do comparative studies of most animals show (speciation type)?
In many animals (mammals), range overlap almost always increases over time, which is consistent with allopatric speciation.
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What is the problem with mitochondrial DNA?
Difficulties in determining close relatives. For example, two very closely related species of chipmunk show little to no divergence with large range overlap based on mtDNA (sympatric). However, when using nuclear DNA, this pattern disappears.
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What animal was different in the comparative studies?
Pocket gophers showed perpetual allopatry (there is no increase in range overlap with time). No matter how long they have diverged, there is no evidence for sympatric speciation.
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What is a possible case of sympatric speciation for a comparative study?
For kangaroo rats, do not tend to see an increase in overlap over time. A point shows that the closest relative have a high degree of overlap.
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Who proposed models of ecological speciation?
Charles Darwin and Ronald Fisher
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When did ecological isolation resurge?
Just recently
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When does ecological isolation take place?
Prezygotic barrier
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What are the three categories of ecological isolation?
Habitat isolation, pollinator isolation, and temporal isolation.
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Habitat isolation
Occurs when spatial separation of species is based on biological differences and reduces gene flow between taxa.
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What does spatial separation of taxa in habitat isolation result from? Doesn't?
From genetic differences in fitness based on habitat use not geographic isolation.
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What is habitat isolation based on?
A species' inability to use another species' environment (can exist in sympatry and adaptations arise). Competition forces species to use different niches when in sympatry and in animals, the ability to seek out habitats for which they are best suited.
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What does habitat isolation limit in animals?
Limits reproductive encounters between conspecifics and heterospecifics.
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What does habitat isolation limit in plants?
Reduces relative amount of heterospecific pollen reaching the stigma.
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Why is hybridization reduced in habitat isolation?
Individuals fail to encounter heterospecifics and sometimes even conspecifics during the breeding season.
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Mircospatial isolation
Members of two species occupy same general area but reproductive encounters reduced by adaptations to/preferences for ecologically different parts of this area.
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Example of microspatial isolation in animals.
Square-tailed drongo (Dicurus ludwigii) and the fork- tailed drongo (D. adsimilis). D. L occupies forest areas and the D. a occupies open areas. May occur closely (50 yards) but won't mate.
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Example of microspatial isolation in plants.
Microspatial isolation of six species of goldenrod along a moisture gradient on a prairie slope. One species prefers more dry soil and the other wet soil, so less likely to encounter during breeding season.
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Macrospatial isolation
Two taxa cannot interbreed because their habitats are allopatric (habitats farly isolated from on another; not a geographic barrier). This isolation is important in parasites and host-specific insects that are dependent on allopatric hosts.
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Example of macrospatial isolation in beetles.
Ophraella chrysomelid beetles are host-specific: * Some allopatric species cannot survive or oviposit (lay eggs) on each other’s hosts - Shows genetic basis for difference
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How can gene flow be reduced by macrospatial and microspatial isolation?
Two species may be largely allopatric but their ranges may overlap in an area containing patches of both habitats.
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Give an example of habitat isolation with macrospatial and microspatial isolation.
Crickets Gryllus pennsylvanicus and G. firmus are largely allopatric. G. pennsylvanicus prefers loamy soils while G. firmus prefers sandy soils. In places where both habitats exist, each species is restricted to its preferred soil type.
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How to demonstrate that microspatial isolation occured?
1. Spatial separation between members of different species is, on average, larger than between members of same species 2. This spatial separation impedes gene flow during the period when both breed simultaneously 3. Reduced gene flow occurs solely through decreasing the chance of cross mating or cross pollination 4. Spatial separation is based on genetic differences between species
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What experiments measure habitat isolation?
Transplant and removal experiments
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Transplant experiment
If cross-pollination or hybridization increases when individuals moved closer together, then they are habitat isolated (physically separation enough to keep them from mating).
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Removal experiment
If removal of one species from a habitat causes another species to expand into that area, microspatial isolation enforced by interspecific competition.
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Why difficult to identify habitat isolation in allopatry?
Allopatry could be caused by habitat isolation or could be unrelated.
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How to measure habitat isolation in allopatry?
Transplant or laboratory experiments
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Transplant or laboratory experiments
If allopatry enforced through habitat isolation, species transplanted to each others’ habitats, or grown under conditions favourable to other species, should not thrive.
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Habitat isolation in flies.
Drosophila sechellia lives only on Seychelle islands * Close relative D. mauritiana on Mauritius * D. sechellia feed and breed only on rotting fruits of Morinda citrifolia, which are toxic to other Drosophila * Differences in D. sechellia oviposition preference, larval survival, and adult survival is genetic * Other Drosophila occupying Seychelles would presumably show complete habitat isolation (would not come in contact with one another because of food preference)
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Is habitat isolation common in the wild?
No, not a lot of data showing that spatial isolation reduces gene flow. For instance, in the goldenrod, even though they are separated in microhabitats, pollinators can move and transfer pollen between.
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What is the strongest evidence for habitat isolation?
Studies in nature combined with lab studies.
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Example of habitat isolation (studied and measured) in Utah.
Two subspecies of sagebrush (Artemisia tridentata tridentata and A. tridentata vaseyana) segregated by altitude in Utah. When the subspecies are transplanted in common garden experiments, each species has higher fitness in soil from its native habitat.
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Example of habitat isolation (studied and measured) in California.
Studies of Horkelia plants in California by Clausen et al. 1940. H. fusca lives above 1400m while H. californica and H. cuneata live along the coast below 1000 m. After one generation in common gardens (green house, which is a controlled habitat), species were transplanted to various habitats. H. fusca showed poor viability and little flowering at low elevations while the low-elevation species did not survive at higher elevations.
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Example of habitat isolation (studied and measured) Michigan.
The apple maggot fly Rhagoletis pomonella feeds and breeds on apples and hawthorn, while R. mendax mates and oviposits only on blueberry. Hybrids are readily produced in field cages but are never found in the field where they co-occur in Michigan.
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Example of habitat isolation (studied and measured) in Nebraska.
Habitat isolation in plains leopard frogs (Rana blairi) and northern leopard frogs (R. pipiens). Where their range overlaps in Nebraska, R. blairi breeds in turbid silty streams while R. pipins prefers clear streams.
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Example of habitat isolation (studied and measured) in birds.
Habitat displacement in birds: Closely-related species occupy more divergent habitat when sympatric – Size-related habitat segregation in Amazonian birds enforced by competition with body size (Robinson and Terborgh 1995) – Congeneric birds (more closely related) are often altudinally segregated in New Guinea (Diamond 1973) * Driven by competition since species expand altitudinal range when congeners are absent
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Example of habitat isolation (studied and measured) in birds.
Evidence in sunfish (Lepomis), rockfish (Sebastes), and damselfish (Stegastes) through artifical removal experiments. Other species take over the habitat when one species is gone, but before, they have own separate habitats
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What do artifical removal experiments show?
Can show whether habitat expansion accompanies the absence of a competitor.
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Importance of habitat isolation in speciation.
Invasion of new environments can produce almost any form of reproductive isolation.
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What study showed that habitat isolation developed before other isolation mechanisms?
Habitat isolation evolved first in greenish warblers, prey size and feeding method appeared later.
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What is the best evidence of habitat isolation come from?
Obligate mutualisms in which plants and insect pollinators have evolved and speciated in tandem (cospeciation).
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How does habitat isolation evolve in allopatry and parapatry?
Initial stages of habitat isolation seen in adaptive differences in habitat use among populations of a single species. Ex. plants that are locally adapted to temperature or soil chemistry or heavy metals near mines
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How does habitat isolation evolve in cospeciation?
Host-specific mating (mate and eat on that host, so fully separated during mating period) Ex. parasitic chewing lice Speciation likely allopatric since they are flightless and their entire life cycle is on a single host.
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Cospeciation in pocket gophers and chewing lice.
Each pocket gopher species have a species of chewing lice. The closest relative of pocket gopher species share a pair of relatives of chewing lice. Show divergence of both at the same time.
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Where else does cospeciation occur? What does this show?
Similar findings in seabirds and their lice, including some sympatric species of seabirds with distinct species of lice. Swiftlets and their lice. That parasite evolution followed host evolution (but why do lice have to diverge).
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How can habitat evolution evolve in sympatry?
If two species diverge in sympatry, habitat isolation through niche differentiation can be a driving force, reduces competition. If sympatry results from secondary contact, reinforcement or competition could lead to habitat isolation. ex. warblers and niche partitioning on trees
418
What is the genetics of habitat isolation?
1. Drosphilia simulans/D. sechellia show habitat isolation based on food preferences: adult tolerance shows 5 or more genes, larval for three or more, adult oviposition showed 2 or more, and rate of egg production showed 4 or more genes 2. Acyrthosiphon pisum pisum alfalfa versus clover race on (p. ahpids) different host plant feeding five genes responsible 3. Mimulus guttatus (monkey flower) populations have one gene in copper tolerance NOT a lot of genes required for habitat isolation
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Pollinator isolation
Gene flow between angiosperms is reduced because different species use different pollinators. - hummingbirds would have flowers with long tubes
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When can you infer pollinator isolation?
If closely-related sympatric species of animal-pollinated plants bloom at the same time and can produce viable hybrids in a greenhouse, but none are found in the wild. Pollinated by different species.
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When is pollinator isolation difficult to determine?
No pollinator specificity as the number of visits by a specific pollinator species is not correlated with pollination ability.
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How to measure pollinator isolation via pollinator specificty?
Dye pollen grains: keep track on plant species where the pollen grains end up. - a study in red and white campions (Silene dioica and S. latifolia) using dyed pollen showed that pollinator isolation is about 0.45
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Examples of pollinator isolation.
Fig and fig wasps, yucca, and yucca moths. One-to-one relationship between host and pollinator species. ex. 750 Ficus fig species pollinated by a similar number of Agaoinidea wasps (each fig species own wasp species for pollination)
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Mutualism
Both individuals benefit in an interaction: plants gets a pollinator and insect gets a breeding site.
425
Examples of pollinator isolation in monkey flowers.
Mimulus lewisii (pink) pollinated by bees and M. cardinalis (red) pollinated by hummingbirds. Strength of isolation based on pollinator visits is 0.98. Can be crossed easily in the greenhouse but only 2 of 2000 seeds collected in the wild produced hybrids.
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What is pollinator isolation produced by?
Pollinator isolation produced by differences in flower color and nectar volume.
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Importance of pollinator isolation in speciation.
Thought to be unimportant but several studies show pollinator isolation. General observation that closely-related sympatric species either use different pollinators or place pollen in different parts of the pollinator’s body.
428
What is pollinator isolation important in?
Two broad comparative studies show strong associations between species diversity and animal pollination or traits associated with animal pollination (in groups where flowers are pollinated by animals there tends to be more species). – Pollinator isolation may have been instrumental in angiosperm speciation
429
How can pollinator isolation evolve?
Easily in allopatry: - Novel pollinators in new areas – Or similar species of pollinators but in different proportions – Extinction of a local plant or pollinator – Change in pollinator preference due to change in visual or chemical environment could be followed by flower evolution to match preference
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Evolution of pollinator isolation example.
Incipient stages of pollinator isolation in the African orchid Satyrium hallackii. Subspecies ocellatum lives in grasslands and is pollinated by hawkmoths. Subpspecies hallackii lives on the coast and is pollinated by carpenter bees. Corresponding changes in flower spur length (the part with nectar in it). Speciation in process.
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Where does weak evidence of of pollinator isolation occur?
In sympatry because how can reproductive isolation evolve when at least one other species – the pollinator – is a key player? (changes in both).
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Genetics of pollinator isolation.
In monkey flowers, genes affect 12 floral traits involved in pollinator isolation (colour, shape, etc.). In Aquilegia species, 1 gene in nectar spurs, 1 gene is shape of spurs, and 1 gene in flower position. NOT much genes needed to control the change.
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Temporal isolation
Gene flow impeded because different species breed at different times.
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Temporal isolation in animals
Different breeding season or spawning season.
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Temporal isolation in plants
Different flowering periods, timing of pollen shedding, timing of ovule receptivity.
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How to detect temporal isolation?
Must show that eliminating the barrier could produce viable and fertile hybrids (breed at the same time). - two types of pink salmon breed every other year - when in the lab, on the same cycle, they can breed together
437
Example of temporal isolation in coral reefs.
Tropical Atlantic corals Montastraea annularis and M. franksi. Release gametes synchronously into the water column over 15-30 minutes. Peak spawning time of two species is 1.5-3 hours apart. Artificial fertilizations produce viable hybrids. Spawning time between M. annularis and M. faveolata is not offset because they are more distantly related and their gametes are incompatible. - important for closely-related species that don't have additional isolating mechanisms
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Importance of temporal isolation in speciation for animals.
General consensus that temporal isolation is not very important in animals. ex. in many birds and mammals, timing of reproduction is tied to food availability and photoperiod – little option for temporal segregation
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Importance of temporal isolation in speciation for plants.
In plants, habitat or pollinator isolation may be more important than temporal isolation.
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When is temporal isolation more common?
In already-isolated species.
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Evolution of temporal isolation in allopatry and paraptry in animals.
In animals, local differences in food availability and temperature (not as common).
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Evolution of temporal isolation in allopatry and paraptry in plants.
In plants, abiotic cues for flowering time such as photoperiod, temperature, and moisture.
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Evolution of temporal isolation in sympatry in plants.
In plants, sympatric species can diverge in flowering time by reproductive character displacement. To reduce competition for pollinators, to prevent hybrid inviability or sterility, or to avoid clogging stigmas with foreign pollen.
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How does temporal isolation evolve in sympatry?
Through reproductive character displacement (any trait that makes plants or animals less likley to mate) and reinforcement (natural selection that trys to enhance mechanisms to keep species apart by selecting against hybrids).
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Evolution of temporal isolation in sympatry in animals.
In animals, reinforcement increases temporal isolation to reduce hybridization or reduce interference between mating calls. ex. spring field cricket and fall field cricket that call and mate at different times OR genus Rana of frogs have the sympatric populations breeding at different times (3)
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How could temporal isolation cause speciation in sympatry?
Instantaneous shift in breeding season. ex. pink salmon that breed in alternate years
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Genetics of temporal isolation.
Lacewing species, two genes are responsible to photoperiods (day lengths) to trigger mating at different times of a year.