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Course Website:

                                                                              http://homepages.ucalgary.ca/~dsikes/courses.htm

                                                                              Check weekly for lecture updates, readings, etc.

                                                                              Course Content:
                                                                                  Theory & Practice of Systematics
                                                                                  Not taxon-specific
Introduction                                                                        25 % alpha taxonomy
to                                                                                  75 % phylogenetics
Biosystematics - Lecture 2




Introduction to Biological Systematics                                       Lecture 2:
                                                                             Introduction to Biological Systematics
Website, Course Outline                                                      Outline: The role and value of Systematics

       Readings (text…, reserve reading room)                                     Taxonomy (! taxonomy) - in decline?
       Midterm & final exam                                                           Describing species
       Project                                                                        Identification, Classification
       “Labs” - Fridays meet in BI 182                                                Collections, Conservation
       Beetle exercise                                                            Phylogenetics (" taxonomy)
       Debates (discussions of readings)                                              Phylogeny
       5 Questions - bonus                                                            Classification (?)
       Open office hours - make appointment                                           Evolutionary processes / patterns
                                                                                      Conservation




Biosystematics                                                               Godfray (2002)
Lecture 1: Introduction to biological systematics (value)

Mayr, E.& P. D. Ashlock (1991) Principles of Systematic Zoology, 2nd           • alpha taxonomy / phylogenetics
Edition, McGraw-Hill, Inc., NY. pp. 1-8. [PDF]
                                                                                   (descriptive taxonomy / phylogenetic tax.)
*Godfray, H. C. J. (2002) Challenges for taxonomy. Nature 417: 17-19.
                                                                                   - suffering from lack of prestige & resources
Flowers et al. (2002) Does the decline in systematic biology matter?
Chapter 4 of report to the House of Lords (UK). Select Committee on
Science & Technology. [webpage]                                                • Solutions?
Lecture 2: Value of biosystematics continued; History of taxonomy

*Gould, S. J. 2000. Linnaeus's Luck? Natural History. Vol 109 iss.7 :
                                                                               • “web monographs” ?
(September): 19-25, 66-76.

Mayr, E.& P. D. Ashlock (1991) Principles of Systematic Zoology, 2nd
                                                                               • unitary taxonomy             ?
Edition, McGraw-Hill, Inc., NY. pp. 9-18. [PDF - see link under lecture 1]




                                                                                                                                   1
Gould - Linnaeus’s Luck                         ! (alpha) taxonomy


  “Descriptive” taxonomy                           “Systematic biology has contracted at
    - Describing                                   British universities to such an extent that it
    - Cataloging                                   may be in danger of extinction as a
    - Classifying                                  sustainable discipline.”
    - Stamp collecting?
                                                   - 1992 the Dainton Report on Systematics
  Hypothetico-deductive taxonomy                           in the UK
    - Descriptions based on theory &
         hypotheses
    - Seek a ‘natural’ classification




! (alpha) taxonomy                              ! (alpha) taxonomy


  Demographic trends in alpha taxonomy:           “If we found these demographic trends in a newly
                                                      discovered species of lemur, we would bring
                                                   specimens into a zoo and start a programme of
  1990 survey                                                      captive breeding.

                                                 But if these trends continue among taxonomists and
  63% of taxonomists > 46 years old               systematists, how soon will it be before we cannot
                                                          recognize a new species of lemur?”
  Only 8% < 35 years old
                                                  Gaston, K. J. and R. M. May. 1992. Taxonomy of
                                                         taxonomists. Nature 356: 281-28




! (alpha) taxonomy                              ! (alpha) taxonomy


Systematics training in universities -         Results of this decline -

     - molecular systematics (your text)            - systematists who can’t do identifications

     - phylogenetics                                - orphaned taxa

     - rarely any training in alpha taxonomy        - fewer people to describe species

     - knowledge & skills are being lost            - lots of new trees, but fewer taxonomic
                                                          changes




                                                                                                       2
“We sit on the brink of    ! (alpha) taxonomy
                                           a crisis”


                                                                     Solutions -

                                                                          - Modernization - new technology

                                                                          - digitization, web-publication, DNA

Not enough trained                                                        - NSF PEET grants
taxonomists to describe
remaining 5-15 million
                                                                              (Partnership for Enhancing Expertise
species - even if we                                                               in Taxonomy)
had the money




                       Biosystematics                                 Classification - usage of term

                                                                      Standard:

             identification
                                       character                      “Activity of grouping entities or phenomena
                                       evolution
                                                                      and giving names to the resulting groups”
             descriptions             phylogeny
                                                                      -Wiley (1981) p. 194

   collections            classification           biogeography




 Classification - usage of term                                       Classification - usage of term

 Actual usage today:                                                  - Not ‘identification’

 “Activity of grouping entities or phenomena                          - Not phylogenetic analysis
 and giving names to the resulting groups”                               - often used as such (e.g. “3 schools of classification”)
                                                                         - implications are wrong

   Phylogenetic analysis                                              - Not a phylogeny
                                                                         - often thought that “tree = classification”
                                  Classification
                                                                         - many students misunderstand this




                                                                                                                                     3
Classification - usage of term                              Example Classification

   So what is it?                                              Phylum Annelida
                                                                    Class Polychaeta
   - is an arrangement of names - names for                                   Family Siboglinidae
         groups (of names…)                                         Class Clitellata
                                                                         Subclass Oligochaeta
        - Linnaeus prepared classifications                              Subclass Hirudinea
                                                                    Class Echiura
        - many modern phylogeneticists do not                  Phylum Sipuncula
             -new trees are not new classifications
             -new classifications can be based on new          We’ll do more on classifications and trees later
                  trees however




Trees                                         Classification   Biosystematics - value
                                              = Names

                                                               1. Provide Classifications for our millions of
                                                                  species

                                                                 Provide natural classifications
                                                                 = reflect evolutionary history
                                                                 = based on sound phylogenetic analyses

                                                                 Natural classifications allow predictions

Giribet et al
Nature 2001




   Biosystematics - value                                      Biosystematics - value


   Allow prediction of attributes of taxa not yet              Bad, “unnatural” classifications
      studied                                                       can be disastrous

        -   medicines (antibiotics, etc.)                      e.g. Gypsy moth - brought to Boston by a
        -   biological control agents                             frenchman, Leopold Trouvelot
        -   predict ecological relationships                      - hoped to start a business in silk
        -   extinct taxa - singing dinosaurs?                     - chose this moth because of its name at
                                                                          that time: Bombyx dispar
                                                                  - oriental silk moth = Bombyx mori




                                                                                                                  4
Biosystematics - value                                 Biosystematics - value


Moth was no good for silk                              (Bad) Taxonomy used for conservation
                                                         assumed 1 species when there were 2+
Was very good at eating native trees
                                                       Resulted in
Escaped & is still a major pest species                  - possible extinction of a subspecies
                                                         - near extinction of a second species
  Current name: Lymantria dispar                                          (extinction of 10 of 40 populations)
  (different superfamily from Bombyx !)
                                                       Described in Nature: 347, 177-179 (1990)




Biosystematics - value                                 Biosystematics - value


2. Infer phylogenies                                   Classified in their own phylum
   - inform our classifications
   - to know what a organism “is” requires             Only in last few decades have we determined
             phylogeny                                   they are Arthropods
       e.g. tongue worms - pentastomids                     specifically Maxillopod Crustaceans!

     bizarre parasites of vertebrates
  - what are they?




Biosystematics - value                                 Evolution of Swim Bladders & Lungs
3. Understanding evolutionary processes               • Both derived from outpocketings of the gut
   a tree is required to study:                       • Both structures hold gas
   - Cospeciation
                                                        – Swim bladders to adjust buoyancy
   - Historical biogeography, phylogeography
                                                        – Lungs for gas exchange
   - Macroevolutionary patterns
      explosive radiations                            • Tetrapods & some fish have lungs
      extinctions                                     • Many fish have swim bladders
   - Trait correlations
     Are warning colors related to the evolution of                     Which is the came first?
          gregariousness?
                                                                        Need a tree…
  - Adaptations




                                                                                                                 5
Biosystematics - value                           Biosystematics - value

Sound natural classifications or phylogenies     Conservation Biology - Biodiversity crisis
  enable biologists to
• intelligently frame their hypotheses

• understand the direction of evolutionary
  change

• know which taxa are appropriate for
  comparative studies
                                                                 Human population




 Biosystematics - value                           Biosystematics - value

Conservation Biology - Biodiversity crisis       Conservation Biology - Biodiversity crisis

 - Massive habitat destruction                    Phylogenies help identify unique lineages

 - ca. 1 species extinction / 20 minutes          e.g. “living fossils”

 - current extinction rates 100 to 1,000 times    e.g. a duckbilled platypus
    greater than “normal”




                                                                                              6
Biosystematics                                                    History of Taxonomy

Systematics "is at the same time the most elementary                                     Father of Biological
and most inclusive part of biology, most elementary                                       Classification
because organisms cannot be discussed or treated in
a scientific way until some taxonomy has been
                                                                                         Used concept of nested sets
achieved, and most inclusive because systematics in
its various branches gathers together, utilizes,
summarizes, and implements, everything that is                                           Named various taxa
known about organisms, whether morphological,
physiological, or ecological.”                                                           e.g. Coleoptera
                                                                                           “Sheath-wing”
Paraphrased from George Gaylord Simpson's book, “Animal Taxonomy”    Aristotle
                                                                                         Oldest scientific names still in
                                                                     384-322 B.C.
                                                                                           use




                          Phylum Annelida
                                                                    History of Taxonomy
                                                                    Botanical taxonomy developed faster than
   Class                  Class                      Class           zoological
   Polychaeta             Clitellata                 Echiura
                           Subclass
                           Oligochaeta
                                         Subclass
                                         Hirudinea
                                                                    Plants used for medicines & foods

                                                                    Theophrastes 371-287 B.C. - classified
                                                                     500 species of plants (e.g. Asparagus)

                           Nested Sets                              Books on plants (herbals) with illustrations,
                                                                     descriptions & names became common




  History of Taxonomy                                               History of Taxonomy
  Herbals copied without improvements                               Phrase names:
   throughout the Middle Ages                                       Joseph Pitton de Tournefort - paved the
                                                                      way for Linnaeus
  Very little progress until the 15th century -
   Gutenberg’s printing press                                       Key work published in 1700 instituted the
                                                                     genus name
  Botanists began using phrase names -
   descriptive names of species                                     e.g. Mentha floribus spicatis, foliis oblongis serratis

                                                                    “mint with flowers in a spike, leaves oblong and
  Latin, usually < 12 words                                           toothed”




                                                                                                                              7
History of Taxonomy                          History of Taxonomy
First Bioinformatics crisis - too many new   The binominal system was actually first
  species & names, chaos reigned              used by Gaspard Bauhin (1623) but
                                              never caught on
Linnaeus instituted a simpler, more
  organized system - binominal               Linnaeus reintroduced & imposed it on the
  nomenclature, 1753, [6,000 plants]           world - chaos was averted

  based on binomen - species name            Zoological nomenclature began with the
  composed of two words - as an               10th edition of Linnaeus’s Systema
  abbreviation for the full phrase names      Naturae (1758) - why this edition?
(note: not binomial nomenclature)




History of Taxonomy                          You should be able to
Linnaeus’s 5 ranks:
                                             Describe the decline of alpha taxonomy &
Kingdom - Animalia                           ideas to reverse it
  Class    - Insecta
                                             Understand the term Classification (not
     Order     - Coleoptera
                                             easy!)
         Genus      - Silpha
               Species - Silpha vespillo     Describe the value of phylogenetics

All animals in 312 genera                    Describe the key people & their influence on
                                             the development of Systematics




                                                                                            8

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Outline taxonomy

  • 1. Course Website: http://homepages.ucalgary.ca/~dsikes/courses.htm Check weekly for lecture updates, readings, etc. Course Content: Theory & Practice of Systematics Not taxon-specific Introduction 25 % alpha taxonomy to 75 % phylogenetics Biosystematics - Lecture 2 Introduction to Biological Systematics Lecture 2: Introduction to Biological Systematics Website, Course Outline Outline: The role and value of Systematics Readings (text…, reserve reading room) Taxonomy (! taxonomy) - in decline? Midterm & final exam Describing species Project Identification, Classification “Labs” - Fridays meet in BI 182 Collections, Conservation Beetle exercise Phylogenetics (" taxonomy) Debates (discussions of readings) Phylogeny 5 Questions - bonus Classification (?) Open office hours - make appointment Evolutionary processes / patterns Conservation Biosystematics Godfray (2002) Lecture 1: Introduction to biological systematics (value) Mayr, E.& P. D. Ashlock (1991) Principles of Systematic Zoology, 2nd • alpha taxonomy / phylogenetics Edition, McGraw-Hill, Inc., NY. pp. 1-8. [PDF] (descriptive taxonomy / phylogenetic tax.) *Godfray, H. C. J. (2002) Challenges for taxonomy. Nature 417: 17-19. - suffering from lack of prestige & resources Flowers et al. (2002) Does the decline in systematic biology matter? Chapter 4 of report to the House of Lords (UK). Select Committee on Science & Technology. [webpage] • Solutions? Lecture 2: Value of biosystematics continued; History of taxonomy *Gould, S. J. 2000. Linnaeus's Luck? Natural History. Vol 109 iss.7 : • “web monographs” ? (September): 19-25, 66-76. Mayr, E.& P. D. Ashlock (1991) Principles of Systematic Zoology, 2nd • unitary taxonomy ? Edition, McGraw-Hill, Inc., NY. pp. 9-18. [PDF - see link under lecture 1] 1
  • 2. Gould - Linnaeus’s Luck ! (alpha) taxonomy “Descriptive” taxonomy “Systematic biology has contracted at - Describing British universities to such an extent that it - Cataloging may be in danger of extinction as a - Classifying sustainable discipline.” - Stamp collecting? - 1992 the Dainton Report on Systematics Hypothetico-deductive taxonomy in the UK - Descriptions based on theory & hypotheses - Seek a ‘natural’ classification ! (alpha) taxonomy ! (alpha) taxonomy Demographic trends in alpha taxonomy: “If we found these demographic trends in a newly discovered species of lemur, we would bring specimens into a zoo and start a programme of 1990 survey captive breeding. But if these trends continue among taxonomists and 63% of taxonomists > 46 years old systematists, how soon will it be before we cannot recognize a new species of lemur?” Only 8% < 35 years old Gaston, K. J. and R. M. May. 1992. Taxonomy of taxonomists. Nature 356: 281-28 ! (alpha) taxonomy ! (alpha) taxonomy Systematics training in universities - Results of this decline - - molecular systematics (your text) - systematists who can’t do identifications - phylogenetics - orphaned taxa - rarely any training in alpha taxonomy - fewer people to describe species - knowledge & skills are being lost - lots of new trees, but fewer taxonomic changes 2
  • 3. “We sit on the brink of ! (alpha) taxonomy a crisis” Solutions - - Modernization - new technology - digitization, web-publication, DNA Not enough trained - NSF PEET grants taxonomists to describe remaining 5-15 million (Partnership for Enhancing Expertise species - even if we in Taxonomy) had the money Biosystematics Classification - usage of term Standard: identification character “Activity of grouping entities or phenomena evolution and giving names to the resulting groups” descriptions phylogeny -Wiley (1981) p. 194 collections classification biogeography Classification - usage of term Classification - usage of term Actual usage today: - Not ‘identification’ “Activity of grouping entities or phenomena - Not phylogenetic analysis and giving names to the resulting groups” - often used as such (e.g. “3 schools of classification”) - implications are wrong Phylogenetic analysis - Not a phylogeny - often thought that “tree = classification” Classification - many students misunderstand this 3
  • 4. Classification - usage of term Example Classification So what is it? Phylum Annelida Class Polychaeta - is an arrangement of names - names for Family Siboglinidae groups (of names…) Class Clitellata Subclass Oligochaeta - Linnaeus prepared classifications Subclass Hirudinea Class Echiura - many modern phylogeneticists do not Phylum Sipuncula -new trees are not new classifications -new classifications can be based on new We’ll do more on classifications and trees later trees however Trees Classification Biosystematics - value = Names 1. Provide Classifications for our millions of species Provide natural classifications = reflect evolutionary history = based on sound phylogenetic analyses Natural classifications allow predictions Giribet et al Nature 2001 Biosystematics - value Biosystematics - value Allow prediction of attributes of taxa not yet Bad, “unnatural” classifications studied can be disastrous - medicines (antibiotics, etc.) e.g. Gypsy moth - brought to Boston by a - biological control agents frenchman, Leopold Trouvelot - predict ecological relationships - hoped to start a business in silk - extinct taxa - singing dinosaurs? - chose this moth because of its name at that time: Bombyx dispar - oriental silk moth = Bombyx mori 4
  • 5. Biosystematics - value Biosystematics - value Moth was no good for silk (Bad) Taxonomy used for conservation assumed 1 species when there were 2+ Was very good at eating native trees Resulted in Escaped & is still a major pest species - possible extinction of a subspecies - near extinction of a second species Current name: Lymantria dispar (extinction of 10 of 40 populations) (different superfamily from Bombyx !) Described in Nature: 347, 177-179 (1990) Biosystematics - value Biosystematics - value 2. Infer phylogenies Classified in their own phylum - inform our classifications - to know what a organism “is” requires Only in last few decades have we determined phylogeny they are Arthropods e.g. tongue worms - pentastomids specifically Maxillopod Crustaceans! bizarre parasites of vertebrates - what are they? Biosystematics - value Evolution of Swim Bladders & Lungs 3. Understanding evolutionary processes • Both derived from outpocketings of the gut a tree is required to study: • Both structures hold gas - Cospeciation – Swim bladders to adjust buoyancy - Historical biogeography, phylogeography – Lungs for gas exchange - Macroevolutionary patterns explosive radiations • Tetrapods & some fish have lungs extinctions • Many fish have swim bladders - Trait correlations Are warning colors related to the evolution of Which is the came first? gregariousness? Need a tree… - Adaptations 5
  • 6. Biosystematics - value Biosystematics - value Sound natural classifications or phylogenies Conservation Biology - Biodiversity crisis enable biologists to • intelligently frame their hypotheses • understand the direction of evolutionary change • know which taxa are appropriate for comparative studies Human population Biosystematics - value Biosystematics - value Conservation Biology - Biodiversity crisis Conservation Biology - Biodiversity crisis - Massive habitat destruction Phylogenies help identify unique lineages - ca. 1 species extinction / 20 minutes e.g. “living fossils” - current extinction rates 100 to 1,000 times e.g. a duckbilled platypus greater than “normal” 6
  • 7. Biosystematics History of Taxonomy Systematics "is at the same time the most elementary Father of Biological and most inclusive part of biology, most elementary Classification because organisms cannot be discussed or treated in a scientific way until some taxonomy has been Used concept of nested sets achieved, and most inclusive because systematics in its various branches gathers together, utilizes, summarizes, and implements, everything that is Named various taxa known about organisms, whether morphological, physiological, or ecological.” e.g. Coleoptera “Sheath-wing” Paraphrased from George Gaylord Simpson's book, “Animal Taxonomy” Aristotle Oldest scientific names still in 384-322 B.C. use Phylum Annelida History of Taxonomy Botanical taxonomy developed faster than Class Class Class zoological Polychaeta Clitellata Echiura Subclass Oligochaeta Subclass Hirudinea Plants used for medicines & foods Theophrastes 371-287 B.C. - classified 500 species of plants (e.g. Asparagus) Nested Sets Books on plants (herbals) with illustrations, descriptions & names became common History of Taxonomy History of Taxonomy Herbals copied without improvements Phrase names: throughout the Middle Ages Joseph Pitton de Tournefort - paved the way for Linnaeus Very little progress until the 15th century - Gutenberg’s printing press Key work published in 1700 instituted the genus name Botanists began using phrase names - descriptive names of species e.g. Mentha floribus spicatis, foliis oblongis serratis “mint with flowers in a spike, leaves oblong and Latin, usually < 12 words toothed” 7
  • 8. History of Taxonomy History of Taxonomy First Bioinformatics crisis - too many new The binominal system was actually first species & names, chaos reigned used by Gaspard Bauhin (1623) but never caught on Linnaeus instituted a simpler, more organized system - binominal Linnaeus reintroduced & imposed it on the nomenclature, 1753, [6,000 plants] world - chaos was averted based on binomen - species name Zoological nomenclature began with the composed of two words - as an 10th edition of Linnaeus’s Systema abbreviation for the full phrase names Naturae (1758) - why this edition? (note: not binomial nomenclature) History of Taxonomy You should be able to Linnaeus’s 5 ranks: Describe the decline of alpha taxonomy & Kingdom - Animalia ideas to reverse it Class - Insecta Understand the term Classification (not Order - Coleoptera easy!) Genus - Silpha Species - Silpha vespillo Describe the value of phylogenetics All animals in 312 genera Describe the key people & their influence on the development of Systematics 8