Phylogeny, Speciation, and Life
Overview of Phylogeny and Speciation
Life originated on Earth over 3.5 billion years ago.
Current estimate of approximately 10 million species on Earth.
Key questions:
How do we define species and how do they evolve?
How can we understand the evolutionary history of Earth?
Understanding achieved through:
Nested patterns of similarities among extant species using phylogenies.
Historical patterns of evolution seen through fossils.
Chapter Interrelationship
Approaching Chapters 24 and 25 simultaneously:
Chapter 24: Discusses how species arise.
Chapter 25: Describes and explains evolutionary relationships between species.
Understanding one chapter requires knowledge of the other.
Preparatory reading for the upcoming class: Finish Chapter 24 and read Bioskills 13; review Chapter 25.1 if time permits.
Speciation and Biological Classification
Chapter 24: Speciation
Opening Roadmap
Learning objectives:
Understanding how speciation creates new branches in the tree of life.
Addressing the following key questions:
How is a species defined?
How do isolation and divergence of populations produce new species?
Allopatry (24.1)
Sympatry (24.2)
Outcomes when isolated populations make contact (24.4)
Binomial Nomenclature
Definition:
The system for naming species, consisting of
Genus name and specific epithet.
Example: Odocoileus virginianus (Virginia deer).
Linnaean System of Classification
Founded by Carl Linnaeus (1707-1778).
Originally phenetic.
System structure bonds life into a nested hierarchical system:
Domain
Kingdom
Phylum
Class
Order
Family
Genus
Species
Mnemonic devices can assist in remembering categories.
Limitations of the Linnaean System
Ranks are categorized arbitrarily – subjective nature of taxon breadth.
Not based on evolutionary history unlike cladistics:
Example: Reptilia (reptiles) and Aves (birds) are classified separately under traditional Linnaean taxonomy; however, evolutionary relationships show that birds are indeed reptiles.
Defining Species
Characteristics of Species
Closed gene pool; alleles shared among members but not usually with other species.
Species can become extinct through genetic divergence lead to new species development.
Local extinction is a concept worth exploring.
Speciation
Definition: Splitting event leading to distinct species stemming from an ancestral species.
Can occur rapidly or gradually.
Involves:
Genetic isolation: Barriers prevent gene flow between populations.
Genetic divergence: Mutation, natural selection, and genetic drift affect each isolated population.
Defining a Species
Taxonomy Questions: Is species definition based on arbitrary naming, or ability to exchange genetic material? The debate includes sexuality as a central aspect.
Biological Species Concept (BSC):
Definition: Groups of interbreeding populations that are reproductively isolated from others.
Notable mechanism of reproductive isolation includes:
Prezygotic Isolation: Preventing mating or fertilization.
Postzygotic Isolation: Hybrid offspring do not survive or are sterile.
Mechanisms of Reproductive Isolation
Type | Description | Example |
|---|---|---|
Prezygotic Isolation | Mating does not occur. | - Temporal: Different mating times (e.g., Bishop and Monterey pines). |
Mating occurs but no zygotes produced. | - Habitat: Different breeding habitats (e.g., mainland and beach mice). | |
Different courtship displays prevent mating. | - Behavioral: Male songbirds sing species-specific songs. | |
Mating fails due to structural incompatibility. | - Mechanical: Lock and key fit in insect genitalia. | |
Gametes incompatible. | - Gametic barrier: Sea urchin sperm and eggs incompatibility. | |
Postzygotic Isolation | Hybrid offspring do not survive. | - Hybrid inviability: Low success rate (e.g., rock dove x ring-necked dove). |
Hybrid offspring are sterile. | - Hybrid sterility: Mules (offspring of horse and donkey). |
BSC Complaints
Challenges in testing BSC.
Not applicable to asexual organisms or extinct species.
Issues arise with ring species and hybridization.
Morphospecies Concept
Definition: Species distinguished by morphological features.
Strengths and Applicability:
Useful in absence of gene flow data.
Applicable to sexual, asexual, and fossil species.
Limitations:
Polymorphisms and cryptic species complicate identification.
Subjectivity in morphological materials leading to inconsistencies.
Phylogenetic Species Concept
Definition: Species identified by evolutionary history; smallest group with a common ancestor.
Key points:
Monophyletic groups include an origin and all its descendants.
Applicable to all population types, including fossils and asexual populations.
May recognize more species compared to other concepts, reflecting life’s diversity.
Application of Species Concepts
Researchers employ all three species concepts to define evolutionarily distinct populations.
Conflicts may arise from differing perspectives on classification.
Additional Definitions of Species
Ecological Species Concept (ESC): Based on ecological niche and resource usage.
Summary Table: Common Species Concepts
Concept | Criterion for Identifying Species | Advantages | Disadvantages |
|---|---|---|---|
Biological Species | Reproductive isolation among population groups | Evolutionary independence evident. | Inapplicable to asexual or extinct species; geographic overlap issue. |
Morphospecies | Morphological distinction among populations | General applicability in lacking genetic data. | Misidentification concerns; subjective morphologies. |
Phylogenetic Species | Monophyletic grouping on phylogenetic tree | Widely applicable and testable criteria. | Currently limited phylogenetic resources. |
Speciation Process
Definition: Development of reproductively isolated populations from genetic divergence.
Mechanisms leading to divergence:
Mutations lead to genetic differences over time.
Speciation occurs when populations can no longer produce viable offspring.
Types of Speciation
Allopatric Speciation: Occurs through geographic separation.
Sympatric Speciation: Occurs without geographic separation.
Mechanisms:
External (e.g., disruptive selection based on ecology).
Internal (e.g., chromosomal mutations).
Allopatric Speciation
Processes:
Dispersal: individuals migrating.
Vicariance: physical barriers such as mountains or rivers.
Example: Genetic divergence through physical separation and habitat change (e.g., Amazon basin's geological division).
Sympatric Speciation
Populations can interbreed but are economically impacted by niche differentiation or species-specific preferences leading to divergence.
Instantaneous speciation through hybridization can lead to distinct reproductive isolation.
Polyploidy, particularly in plants, plays a significant role in sympatric speciation.
Outcomes of Secondary Contact
Processes Upon Fusion of Isolated Populations:
Fusion leading to interbreeding.
Extinction of one species.
Reinforcement of divergence through natural selection against hybrids.
Hybrid zone formations with geographic movements.
Formation of new species leading to new ecological niches.
Large-Scale Patterns in Life’s History
Adaptive Radiation
Definition: Rapid emergence of many distinct species from a single ancestor due to diverse adaptations.
Examples:
Hawaiian silverswords evolved from tarweed, showing monophyly, rapid speciation, and ecological diversity.
Analysis of factors prompting adaptive radiation includes extrinsic and intrinsic triggers.
Factors Triggering Adaptive Radiations
Extrinsic Factors: Favorable new environmental conditions.
Intrinsic Factors: Evolution of crucial morphological, physiological, or behavioral traits (such as the development of flowers or birds' flight adaptations).
Ecological Opportunity and its Role
Defined as the availability of various resources for exploitation and diversification.
As observed in Hawaiian islands where flowering plants were few, leading to ecological diversification by silverswords.
Conclusion: The Case of Galapagos Finches
Overview of natural selection's role in species development among Galapagos finches, with multiple case studies illustrating allopatric speciation and adaptive radiation principles culminating in notable evolutionary patterns and adaptations.