Lecture 11.
The
Contributions of
Systematics To
Evolutionary Biology
Darius Noel C. Minoza
Department of Biological Sciences
College of Arts and Sciences
Visayas State University
Introduction
•Common Misconception
  •Classification of the living world is virtually
   complete
  •We can use any of a range of books to identify
   and classify any common organism
Introduction
•However,
  •Careful comparison of
   several texts reveals
   differences in treatments
  •Examples
    • Family placement of a species (Durian:
      Bombacaceae, Malvaceae)
Introduction
•Hence,
  •Taxonomists have been accused of:
   ✔Pervert desire to confuse things
   ✔Unscientific
   ✔Subjective
   ✔Lack of better things to do
Introduction
•Actually,
  •Reflects ongoing differences in opinions
   concerning the most appropriate
   classification
  •Continuing acquisition of additional
   knowledge
  •Active intellectual exchange and healthy
   scientific debate
So what is TAXONOMY?
•Greek taxis (arrangement), nomos (law)
•Theory and practice of classifying
 organisms (Mayr et al.)
•Recognizing, naming and classifying
 organisms (Gullan & Cranston, 2000)
•How does it differ from systematics?
and what about
SYSTEMATICS?
•Latinized Greek word systema as applied to
 systems of classification developed by early
 naturalists especially Linnaeus (1735)
and what about SYSTEMATICS
•Scientific study of the kinds and diversity of
 organisms and of any and all relationships
 among them (Simpson, 1961)
•Study of the kinds and diversity of
 organisms and their interrelationships
 (Gullan & Cranston, 2000)
•Science of the diversity of organisms (Mayr,
 1969)
Important aspects of the
definitions
•Organism
•Science/Scientific study
•Kinds
•Diversity
•Relationships
•Theory
•Practice
•Classifying
Classification
•Ordering of organisms into groups or
 sets on the basis of relationships
•Ordering of populations and groups of
 populations at all levels by inductive
 procedures
•Most important aspects:
  •Grouping
  •Ranking
Identification
•Placing of individuals by deductive
 procedures into previously established
 classes
Nomenclature
•Application of distinctive names to each
 of the groups recognized in a
 classification system
Taxon
•A group of real organisms recognized as
 a formal unit at any level of a hierarchic
 classification
•”A taxonomic group of any rank that is
 sufficiently distinct to be worthy of being
 assigned to a definite category
Taxon: Two Important
Aspects
•Concrete zoological objects (organisms)
•Formally recognized by a taxonomist
•NOTE: races, biotypes, isolates, forms
 become taxa only when they are
 formally recognized as species or
 subspecies
Taxonomic Categories
•Original Linnean hierarchy
•Only five within the animal kingdom
  •Classis
  •Ordo
  •Genus
  •Species
  •Varietas (Linnaeus’ optional category for various
   types of infraspecific variants; now discarded in
   zoology)
Taxonomic Categories
•Linnaean hierarchy   •Present standard
  • Regnum              • Kingdom
                        • Phylum (=Division)
  • Classis             • Class
  • Ordo                • Order
                        • Family
  • Genus               • Genus
  • Species             • Species
  • Varietas
Additional Taxonomic
Categories
•Empire
•Domain
•Tribe (Between family and genus)
•Cohort (Between class and order in
 some vertebrate groups but between
 order and family among mites)
More detailed taxonomic
categories
•Super
•Sub
•Infra
•Informal categories
   •Series
   •Clade (Cladus)
   •League (Legio)
   •Section (Sectio)
More detailed taxonomic
categories
•Below the species, the only accepted
 category in zoology is the subspecies
•In botany, additional infraspecific
 categories aside from subspecies are:
  •Variety (including cultivar, pathovar)
  •Forma
Acceptable Abbreviations
 •Genus – gen. or g.
 •Species (singular) – sp.
 •Species (plural) – spp.
 •Subspecies (singular) – ssp. or subsp.
 •Subspecies (plural) – sspp. or subspp. for
  plants
 •Variety – var. (cultivar – cv.; pathovar – pv.)
 •Forma – f.
Classification of the Oriental
Weaver Ant
Kingdom Animalia
  Subkingdom
    Phylum Arthropoda
      Subphylum Mandibulata
         Superclass Hexapoda
           Class Insecta
             Subclass Pterygota
                Infraclass Neuroptera
Classification of the Oriental
Weaver Ant
Supercohort
  Cohort
    Infracohort
       Superoder Hymenopteroidea
         Order Hymenoptera
            Suborder Apocrita
              Infraorder
                 Epifamily
Classification of the Oriental
Weaver Ant
Superfamily Vespoidea
  Family Formicidae
    Subfamily Formicinae
       Infrafamily
          Supertribe
             Tribe Formicini
                Subtribe Formicina
                  Infratribe
Classification of the Oriental
Weaver Ant
Genus Oecophylla
  Subgenus Oecophylla
      Species smaragdina
         Subspecies smaragdina
Full scientific name
  Oecophylla (Oecophylla) smaragdina smaragdina
(Fabricius)
Phylogenetics
•the unravelling of evolutionary history
•controversial area of biology
•morphology – provided much of the data upon
 which relationships of organisms were
 reconstructed
•some of the “ambiguity” and lack of clarity
 regarding phylogeny of many groups
   •blamed on inherent deficiencies of phylogenetic
    information in morphological characters
Phylogenetics
•Other data sources explored
•Cytogenetic – mainly chromosome numbers
•Allozymes, isozymes (relies on differences in
 electrophoretic mobility of proteins),
 biochemical
•Nucleic acid sequence (DNA, RNA), data
 (molecular)
Phylogenetics
•Increased use of sequence data (molecular)
•Attempts to resolve some unanswered
 questions (esp. higher relationships of
 insects etc.)
•Appropriate choice of taxa and genes,
 molecules are providing resolution of some
 phylogenetic questions that morphology has
 been unable to answer
Phylogenetics
 •Preferred approach to estimating
  phylogenies:
1. Holistic
   -Use data from as many sources as
    possible
   -Be constantly aware that:
     -Not all similarities are equally informative in
      showing phylogenetic patterns
Phylogenetic methods
•Methods that attempt to recover or
 reconstruct the pattern produced by
 phylogenetic history
•Rely on observations of living organisms and
 fossils
Phylogenetic Methods
•Phenetics
•Cladistics
•Evolutionary Systematics
Phenetics
•Also known as traditional
 taxonomy
•Quantitative or numerical
 method for distinguishing
 species based on
 morphological grounds
•Based on overall
 morphology, especially
 quantitative data from
 morphometric studies
Examples (Bivariate plot)
Examples (GM- Geometric
morphometrics)
Cladistics
•Quantitative method that
 uses shared character to
 create hypothesis
•Rely on assumptions
 about ancestral
 relationships as well as
 current data
Consider these taxa
•Monophyletic
-Descendants of a
single common
ancestor
Consider these taxa
•Paraphyletic
-Do not include all
descendants of a
single common
ancestor
Consider these taxa
•Polyphyletic
-Do not share
common
ancestor and have
multiple origins
Evolutionary Systematics
•Also referred to as Classical Evolutionary
 Systematics/Taxonomy, Evolutionary
 Taxonomy
•Uses estimates of derived similarity
•UNLIKE Cladistics
 ✔Estimates of the amount of evolutionary change are
  included with the branching pattern in order to
  produce a classification
Evolutionary Systematics
•Emphasizes distinctness
•Grants higher taxonomic status to taxa
 separated by “gaps”
•May be due to accelerated morphological
 innovation in a lineage and/or extinction of
 intermediate linking forms
Phylogenetic Methods
•The case of the ants
  •Highly specialized
  •Many unique features
  •Looks very different from nearest relatives
    •Frequently given superfamily rank (Formicoidea)
     within Hymenoptera (evolutionary taxonomy)
Phylogenetic Methods
•The case of the ants
  •Phylogenetic study indicates
    •Family Formicidae in the Superfamily Vespoidea (a
     group of solitary,semi-social and social wasps
Phylogenetics
•Paraphyletic groups – avoid if possible
 because their only defining features are
 ancestral characters shared with other
 indirect relatives
•Thus, absence of wings in paraphyletic
 apterygotes is an ancestral feature shared
 by many other invertebrates
Phylogenetics
•Polyphyletic groups – their mixed ancestry
 means that such groups are biologically
 uninformative
•Such taxa are artificial and should never be
 included in any classification
Phylogenetics
•Current classification of most organisms
 1. Mix all three practices
 2. Most orders based on groups (taxa) with
    most distinctive morphology
 3. Such morphologically distinct groups are not
    necessarily monophyletic
Phylogenetics
•Current classification of most organisms
•Some are almost certainly paraphyletic
•However, it is unlikely that any higher-level
 groups are polyphyletic
Phylogenetics
•Colloquial terms
•Flies – cover disparate groupings from
 mayflies (Ephemeroptera) to true flies
 (Diptera)
•Bugs - ! ! !
Phylogenetics
•Refinements continue
•Classification: should reflect our developing
 understanding of phylogeny
•Current classification – combination of
 traditional views variably reconciles with
 phylogeny
Three Schools of Taxonomy
•Consider how the three schools of taxonomy
 will classify groups of animals that were first
 taught to us as kids -
•Four-legged (tetrapods), amniotic
 vertebrates (mammals, birds, reptiles)
Classification of Amniote
Tetrapods (Evolutionary Taxonomy)
  Class Mammalia
  Class Reptilia
    Subclass Testudines (turtles)
    Subclass Squamata (lizards, snakes, crocodiles)
  Class Aves
Classification of Amniote
Tetrapods (Phenetics)
•Class Mammalia
•Class Reptilia
 •Subclass Testudines
 •Subclass Squamata
•Class Aves
Classification of Amniote
Tetrapods (Cladistics)
•Class Mammalia
•Class Reptilia
 •Subclass Testudines
 •Subclass Sauria
   •Infraclass Squamata (lizards, snakes)
   •Infraclass Archosauria (crocodiles, birds)
     • Order Crocodilia
     • Order Aves
                       Naming based on current data
Naming based on past data
                         Reptiles
                                                      Sauropsida
Three Schools of
Taxonomy
•Note similar examples among insects and
 many other groups of organisms
•More expected to be revealed by
 •Molecular data
 •More detailed morphological data
  (ultrastructure)
Trees: What they are
•Cladogram – a tree-like diagram showing
 hierarchic relationships, to depict relative
 recency of such relationships, without
 connoting degree of difference
•Phylogenetic trees or cladograms are
 diagrammatic representations of
 phylogenetic relationships
Cladogram
Trees: What they are
•Dendrogram – a tree-like diagram intending
 to show both relationships (branching or
 “cladogenesis”) and degree of difference
 (divergence or “anagenesis”) (evolutionary
 taxonomy)
Trees: What they are
•Phenogram – a diagram representing
 hierarchic relationships obtained by applying
 phenetic techniques, whereby data are
 clustered at levels (phenetics)
Trees: What they are
•Cladograms result from analyses of sets of
 data – morphological and or molecular
•Can only be as good as the available data
 could support
•Therefore, cladograms are diagrams
 depicting hypotheses of relationships
Trees: terminologies
•Character – a feature showing a variation(s)
 that define a group
•Character-state – one of the various
 conditions of a feature (character) obtained
 across a group of taxa
•Character-state tree – graphic
 representation of the coding of a character
Trees: terminologies
•Apomorphy- advanced or derived character;
 a group-defining character (adj. apomorphic)
•Pleisomorphy- “primitive” character;
 ancestral character; not group defining at the
 level at which it is being observed; the
 quality being primitive (adj. pleisomorphic)
Trees: terminologies
•Synapomorphy- shared derived character
•Symplesiomorphy- shared primitive
 character
Figure 4.5, pg. 116
Trees: What they are and
how they are constructed
 •Parsimony
–seeks the simplest explanation that requires the
least amount of change (fewest steps)
 •Most parsimonious tree (MPT) – for a given
  data set, tree(s) of minimum length as
  computed with parsimony as main criterion
 •Consensus tree – depicts “compromise” of all
  MPTs
Parsimony
Parsimony
Parsimony
Parsimony
Parsimony
Trees: What they are and
how they are constructed
•Data matrix- information in tabular form on
 characters (columns) for a set of taxa (rows)
•Terminal taxon – taxon for which data are
 actually coded in a cladistic analysis
•Hypothetical ancestor – node in a
 cladogram representing the deduced set of
 attributes for two or more terminal taxa
Data Matrix
Data Matrix
Trees: How they are
constructed
  •Selection of ingroup and outgroup taxa
1. For rooting of trees
2. Character polarity
  i. Character analysis
 ii. Construction of data matrix
iii. Coding methods
iv. For molecular data – alignment of nucleic
     acid sequence
Trees: How many trees?
•Number of possible rooted bifurcating trees
 for 1 to 22 taxa
-Can range from 1 to 13,113,070,
 457,687,988,603,440,625
-Now possible through available
 software/programs
Trees
Trees
Trees
Types of Phylogenetic
Analysis Methods
1. Phenetic
   i.   UPGMA (Demonstration)
  ii.   Fitch-Margoliash method
 iii.   Neighbor joining method
2. Cladistic
 iv.    Parsimony
  v.    Maximum Likelihood
Distance Methods
•Optimal tree is generated by calculating an
 estimate of the evolutionary distance between all
 pairs of sequences first
Example
Example (A and B)
Example (A and C)
Example (A and D)
Example (A and E)
Example (A and F)
Example
Example
How to prove?
Let us try!
•Measure
distance
between
branch tips
Let us try!
•Count the
number
of nodes
between
species
Let us try!
•Compare
time to a
common
ancestor
Let us try!
•Compare
number of
shared
monophyletic
groups
Assessing Confidence Level
of Phylogenetic Trees
•Bootstrap Method
   •Randomly generates new data sets from
    the original set (1000 replicates is most
    common)
   •Computes the number of times that a
    particular grouping (or branch) appeared
    in the tree.
Sample Studies on
Phylogenetic Analysis
Sample Studies in
Phylogenetic Analysis
Samples studies on
Phylogenetic Analysis
Workflow in Phylogenetic
Analysis