Phylogenetics (long)
Evolutionary History of Speciation
Evolutionary history is composed of a series of speciation events.
This history of speciation involves two fundamental elements: - Anagenesis: Character modification. - Cladogenesis: Lineage splitting.
As long as anagenesis keeps pace with cladogenesis, this history of speciation can be recovered.
The Process of Science
Science as a process is centered around the generation of hypotheses and hypothesis testing.
Generating hypotheses about the sequence of speciation leading to biodiversity (both extant and extinct) is the role of the systematic biologist.
Systematics is crucial in evolutionary investigation; it is impossible to engage in evolutionary hypothesis testing without reference to a phylogenetic hypothesis.
Major Divisions in Biology
According to Nelson (1970), biology can be divided into: - General Biology: Concerned with process and mechanism; focuses on the best-suited groups to examine these processes. - Comparative Biology: Concerned with diversity and the causal explanation of that diversity, focusing on: 1. Distributions of characters among taxa. 2. Origin of characters. 3. Reasons for the retention of particular characters.
Systematics and Taxonomy
Systematics: Studies organismic diversity in relation to relationships among populations, species, or higher taxa (Wiley, 1981). All comparative data may be useful in systematic studies.
Taxonomy is related to systematics but is not synonymous. Taxonomy (from Greek, meaning arrangement and law) focuses on naming and classification.
Phylogenetic Systematics
Phylogenetics: A branch of systematics aimed at recovering phylogenetic relationships and producing classifications to reflect these relationships. - The goals include recovery of speciation history on this planet.
Original theory of phylogenetics published by Willi Hennig in 1950, outlining five basic ideas: 1. Cohesion in living and extinct organisms is through genealogical ("blood") relationships. 2. Relationships exist among individuals within populations, between populations, and among species. 3. Other relationships (phenotypic, genetic) correlate with genealogical descent and modification (evolution). 4. Genealogical relationships can be discovered through character examination. 5. The best classification of organisms reflects their genealogical relationships.
Essential Terminology in Phylogenetics
Taxon: A grouping of organisms with a proper name.
Natural Taxon: A group that reflects a unique history of descent (e.g., Angiospermae).
Species: Lineages independent of others; the highest level of taxonomic organization.
Monophyletic Group: Contains an ancestor and all descendants (a clade).
Paraphyletic Group: Contains an ancestor and some, but not all descendants.
Polyphyletic Group: Lacks the most recent common ancestor of its members.
Sister Group: Closest relative of a taxon.
Out-Group: A species used to determine character states (apomorphic vs. plesiomorphic).
Character: An observable feature of an organism (e.g., flower color).
Evolutionary Novelty and Homology
Evolutionary Novelty: Inherited changes in character states (e.g., the change from hair to feathers). - Most novelties undergo polymorphism before becoming fixed.
Homology: Features traced back to a common ancestor versus analogy (similarities without a common ancestor). - Apomorphy: A derived character. - Plesiomorphy: A primitive character. - Synapomorphy: A shared derived character. - Symplesiomorphy: A shared primitive character.
Goals of Taxonomy/Systematics
Recognize, describe, and classify new species.
Examine population structure.
Reconstruct genealogical history of lineages.
Investigate speciation processes and components.
Explore biogeography.
Historical Overview of Taxonomy
Pre-Darwinian Taxonomy
Aristotle: Credited with writing down biological classifications; believed in perfect forms without appreciation for variation. Recognized major groups (e.g., birds, insects). - Emphasized multiple attributes for classification.
Post-Darwinian Taxonomy
Shift from essentialism to recognition of species as mutable entities.
Systematics gained prominence at the expense of taxonomy.
Linnaeus: Developed hierarchical classification and binomial nomenclature, categorizing organisms based on criteria like morphology.
Linnaean Hierarchy
Finalized in 1758; categorizes organisms into: - Two Kingdoms, Five Categories: Class, Order, Genus, Species, Variety. - In zoology: Kingdom, Phylum, Class, Order, Family, Genus, Species. - In botany: Kingdom, Division, Class, Order, Family, Genus, Species.
Schools of Systematic Thought
Phenetics: Grouping based on overall similarity.
Traditional Taxonomy: Emphasizes empirical classification without philosophical background.
Cladistic Taxonomy: Relies on synapomorphies for classification.
Evolutionary Systematics: Integrates Darwinian principles with classification based on genealogical patterns and synapomorphies.
Evolutionary Systematics Goals
Provide classifications reflecting evolutionary phenomena.
Justify the naming and ranking of paraphyletic groups based on morphological gaps, species richness, and adaptive zones.
Phenetics vs. Evolutionary Systematics
Phenetics aims for objectivity, applying equal weight to characters without initial assumptions about their significance.
Character Weighting: Used in evolutionary systematics to emphasize certain characters over others, seen as subjective in contrast to phenetic methods.
Phylogenetics
Analyzes relationships based on shared derived characters (synapomorphies).
The importance of Out-group Comparison for determining character polarity. - Characters must be coded and homologous to derive phylogenetic hypotheses.
Taxonomic Character Types
Structural: Similar in structure, possibly homologous.
Functional: Similar in function, but not homologous (e.g., bird and insect wings).
Phylogenetic: Presumed homologous characters regardless of morphology or function.
Types of Biochemical Characters
Isoenzymes: Variations in proteins indicating genetic diversity.
DNA Sequencing: Techniques including PCR and automated sequencing for analyzing genetic variability.
Species Concepts
Diverse concepts exist for species, including:
Typological: Idea of immutable species types.
Nominalistic: Species as constructs without objective reality.
Biological: Interbreeding populations that are reproductively isolated.
Evolutionary: Lineages that maintain identity and unique evolutionary tendencies.
Phylogenetic: Smallest recognizably diagnosable clusters reflecting ancestry
Modes of Speciation
Sympatric: Populations occupy the same area.
Allopatric: Populations are geographically isolated.
Parapatric: Populations with adjacent ranges that do not overlap.
Stasipatric: Chromosomal mutations lead to speciation.
Cryptic Species: Species that do not display distinct morphological features but are biologically distinct.
Detailed Speciation Models
Vicariance Speciation: Physical separation leads to independent lineages.
Peripheral Isolates: New species emerging in marginal habitats.
Asexual Species Model: Evolutionary stasis provides species cohesion without gene flow.