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:

    1. Nested patterns of similarities among extant species using phylogenies.

    2. 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:

    1. Domain

    2. Kingdom

    3. Phylum

    4. Class

    5. Order

    6. Family

    7. Genus

    8. 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:

    1. Prezygotic Isolation: Preventing mating or fertilization.

    2. 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.