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Comparative Anatomy                                                         page 1



                      Vertebrate Phylogeny
You need to know the phylogeny on the following page extremely well. It is crucial to
understanding the evolution of various anatomical systems. Don't just memorize the
tree. Work with it and understand how it is put together.

                                  Classification

T     he traditional classification (in the Kardong textbook, Appendix D, pp. 696-698) of
      the living vertebrates places them in superclass Agnatha, and classes
      Chondrichthyes, Osteichthyes, Amphibia, Reptilia, Aves, and Mammalia. Of these
only the Chondrichthyes, Aves, and Mammalia are monophyletic. The others
(Agnatha, Osteichthyes, Amphibia, Reptilia) are paraphyletic.

In this course, we either redefine the content of these taxa so that they are
monophyletic, or we abandon usage of the names. Some taxa whose names we have
redefined are Reptilia, Osteichthyes, Sarcopterygii, Amphibia, and Synapsida. These
are redefined to be monophyletic by having the ancestor include all, rather than only
some, of its descendants. For example, Reptilia now includes all of its descendants,
including Aves (birds). Amphibia, which formerly included all early Tetrapoda, but
excluded reptiles, birds, and mammals, now only includes the ancestor of living
amphibians and all of the descendants of that ancestor. See the figures on the next
page.

Taxon names that are capitalized are proper nouns, and represent monophyletic taxa. If
names that refer to paraphyletic groups are used, they are used informally and are not
capitalized, or else put in quotes (e.g., agnathans or quot;Agnathaquot;). Two of the pages
below indicate several commonly used paraphyletic groups.

Don't even worry about learning the classifications given in your textbook on pp. 696
and in the lab book on pp. 32-33.
Comparative Anatomy                                                 page 2

                                                             Hemichordata
                                                             Urochordata
                                                             Cephalochordata
   Chordata
   Euchordata                                                Myxinoidea
                                                             Petromyzontidae
          Craniata
                                           Chondrichthyes    Elasmobranchii
             Vertebrata
                                                             Holocephali
                                                             Brachiopterygii
                                   Actinopterygii
                                                             Chondrostei
              Gnathostomata                                  Lepisosteus
                                                             Amia
                                            Neopterygii
                                                             Teleostei
                  Osteichthyes
                                                             Actinistia
                                                             Dipnoi
                                                             Gymnophiona
                     Sarcopterygii            Amphibia
                                                             Caudata
                            Choanata                         Anura
                                                             Monotremata
                                             Mammalia
                                                             Eutheria
                              Tetrapoda
                                                             Marsupialia
                                                   Theria
                                                             Testudinata
                                   Amniota                   Sphenodontia
                                                             Lizards
                                                             Lizards
                                      Reptilia
                                                             Lizards
                                                             Amphisbaenia
                                          Sauria
                                                             Ophidia
                                                      Squamata
                                                   Lepidosauria
                                                             Aves
                                                             Crocodylia
                                             Archosauria



 Fall 2001
Comparative Anatomy                                                                     page 3


Phylogenetic Taxonomy of Living Vertebrates
A phylogenetic taxonomy of living vertebrates is given below as an indented list; no
fossils are included (yet). A quot;phylogeneticquot; taxonomy means that the tree can be derived
exactly from this list, and vice-versa. The placement of certain fossils will be discussed
as they become relevant to the course. Notice that no ranks (e.g., order, subclass) are
used. The cladistic classification (p. 699) in Kardong is an attempt to do the same thing,
but it is not very well done, so please ignore it.


Craniata
        Myxinoidea
        Vertebrata
                Petromyzontia
                Gnathostomata
                       Chondrichthyes
                               Holocephali
                               Elasmobranchii
                       Osteichthyes
                               Actinopterygii
                                       Brachiopterygii
                                       Unnamed taxon
                                                Chondrostei
                                                Neopterygii
                                                       Lepisosteus
                                                       Unnamed taxon
                                                               Amia
                                                               Teleostei
                               Sarcopterygii
                                       Actinistia
                                       Choanata
                                                Dipnoi
                                                Tetrapoda
                                                       Amphibia
                                                               Gymnophiona
                                                               Batrachia
                                                                        Anura
                                                                        Urodela
                                                       Amniota
                                                               Mammalia
                                                                        Monotremata
                                                                        Theria
                                                                                Marsupialia
                                                                                Eutheria
                                                               Reptilia
                                                                        Testudines
                                                                                Sauria
                                                                                Lepidosauria
                                                                                        Sphenodontia
                                                                                        Squamata
                                                                                Archosauria
                                                                                        Crocodylia
                                                                                        Aves
Comparative Anatomy                                            page 4




                                          Hemichordata
     Nonmonophyletic groups, I
                                          Urochordata

                                          Cephalochordata

                                          Myxinoidea

                                          Petromyzontia
 quot;Invertebrataquot;
                                          Chondrichthyes

                                          Actinopterygii

                                          Actinistia

                                          Dipnoi

                                          Amphibia

                                          Mammalia
             quot;Piscesquot;

                                          Testudinata
          quot;Crossopterygiiquot;
                                                           quot;Homothermiaquot;
                                          Lepidosauria

                             quot;Reptiliaquot;   Crocodylia

                                          Aves
Comparative Anatomy                                      page 5



                                          Hemichordata
     Nonmonophyletic groups, II
                                          Urochordata

                                          Cephalochordata

                                          Myxinoidea

                                          Petromyzontia

                                          Chondrichthyes

                                          Actinopterygii

                                          Actinistia

                                          Dipnoi

       quot;Agnathaquot;                          Amphibia

                                          Mammalia
             quot;Osteichthyesquot;
               quot;Sarcopterygiiquot;            Testudinata
                      quot;Amphibiaquot;
                                          Lepidosauria
                quot;Labyrinthodontiaquot;
                            quot;Synapsidaquot;
                                          Crocodylia

                                          Aves
Comparative Anatomy                                                                       page 6

                          Numbers of Living Species of Chordates
There are about 50,000 species of living vertebrates; this estimate varies, primarily
depending on how many species of teleost fishes are recognized.

Clade Name                             Species     Common Name
Hemichordata                           100         acorn worms, pterobranchs
Urochordata (tunicates)                2000        sea squirts, larvaceans
Cephalochordata                        45          amphioxus or lancelets

Myxinoidea                             30          hagfish
Petromyzontia                          40          lampreys

Chondrichthyes
       Elasmobranchii                  670         sharks, skates, rays
       Holocephali                     30          ratfish or chimaeras

Actinopterygii
       Brachiopterygii                 13          reedfish or bichirs
       Chondrostei                     22          sturgeon, paddlefish
       Lepistosteus                    7           gar
       Amia                            1           bowfin or choupique
       Teleostei                       25000       tarpon, herring, perch, catfish, minnows,
                                                   bass, etc.

Actinistia                             1 (maybe    coelacanth (Latimeria)
                                       2)

Dipnoi                                 6           lungfish

Amphibia
      Gymnophiona                      *160        caecilians
      Caudata                          *350        salamanders
      Anura                            5000        frogs

Mammalia
     Monotremata                       6           platypus, echidna
     Marsupialia                       240         opposum, kangaroos, koalas, wombats,
                                                   etc.
         Eutheria                      4300        mice, elephants, horses, whales, bats, cats,
                                                   etc.

Testudinata                            +230        turtles, tortoises, terrapins

Lepidosauria
       Sphenodontia                    2           tuatara (Sphenodon)
       Squamata
              lizards                  +3000       lizards
              Ophidia                  +2330       snakes
              Amphisbaenia             +140        amphisbaenians

Aves                                   8700        birds
Crocodylia                             +21         alligators, crocodiles, gavials
Numbers from Walker, 1987 and Wake, 1979, except as noted.
*Duellman and Trueb, 1986
+Dowling and Duellman, 1978
Comparative Anatomy                                                           page 7


                                Geological Time Scale
If the entire geological history of earth were set to a one-year scale, the earth formed on
1 January, life appeared on 4 April, and Homo sapiens did not appear until very late on
New Year's Eve.

One-Year Scale                 Real Time                      Event
1 Jan                          4.6 by                         Origin of the earth
4 Apr                          3.8 by                         Life appears (bacteria)
9 Nov                          650 my                         Marine invertebrates
                                                              appear
20 Nov                         520 my                         Vertebrata
29 Nov                         415 my                         Ostracoderms
30 Nov                         410 my                         Gnathostomata
                                                              (acanthodians)
                                                              Actinopterygii
2 Dec                          380 my                         Chondrichthyes
3 Dec                          360 my                         Tetrapoda
7 Dec                          310 my                         Amniota
16 Dec                         190 my                         Mammalia
20 Dec                         150 my                         Aves
26 Dec                         65 my                          Primates
29 Dec                         30 my                          Anthropoidea (monkeys
                                                              and apes)
11:45 pm 31 Dec                125-250,000 yr                 Homo sapiens

Check out these web pages on the Tree of Life (ag.arizona.edu/tree/)
  • Vertebrates
  • Gnathostomata
  • Terrestrial vertebrates (Tetrapods)

Updated by David Cannatella, September 18, 2001

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Vertphylogeny

  • 1. Comparative Anatomy page 1 Vertebrate Phylogeny You need to know the phylogeny on the following page extremely well. It is crucial to understanding the evolution of various anatomical systems. Don't just memorize the tree. Work with it and understand how it is put together. Classification T he traditional classification (in the Kardong textbook, Appendix D, pp. 696-698) of the living vertebrates places them in superclass Agnatha, and classes Chondrichthyes, Osteichthyes, Amphibia, Reptilia, Aves, and Mammalia. Of these only the Chondrichthyes, Aves, and Mammalia are monophyletic. The others (Agnatha, Osteichthyes, Amphibia, Reptilia) are paraphyletic. In this course, we either redefine the content of these taxa so that they are monophyletic, or we abandon usage of the names. Some taxa whose names we have redefined are Reptilia, Osteichthyes, Sarcopterygii, Amphibia, and Synapsida. These are redefined to be monophyletic by having the ancestor include all, rather than only some, of its descendants. For example, Reptilia now includes all of its descendants, including Aves (birds). Amphibia, which formerly included all early Tetrapoda, but excluded reptiles, birds, and mammals, now only includes the ancestor of living amphibians and all of the descendants of that ancestor. See the figures on the next page. Taxon names that are capitalized are proper nouns, and represent monophyletic taxa. If names that refer to paraphyletic groups are used, they are used informally and are not capitalized, or else put in quotes (e.g., agnathans or quot;Agnathaquot;). Two of the pages below indicate several commonly used paraphyletic groups. Don't even worry about learning the classifications given in your textbook on pp. 696 and in the lab book on pp. 32-33.
  • 2. Comparative Anatomy page 2 Hemichordata Urochordata Cephalochordata Chordata Euchordata Myxinoidea Petromyzontidae Craniata Chondrichthyes Elasmobranchii Vertebrata Holocephali Brachiopterygii Actinopterygii Chondrostei Gnathostomata Lepisosteus Amia Neopterygii Teleostei Osteichthyes Actinistia Dipnoi Gymnophiona Sarcopterygii Amphibia Caudata Choanata Anura Monotremata Mammalia Eutheria Tetrapoda Marsupialia Theria Testudinata Amniota Sphenodontia Lizards Lizards Reptilia Lizards Amphisbaenia Sauria Ophidia Squamata Lepidosauria Aves Crocodylia Archosauria Fall 2001
  • 3. Comparative Anatomy page 3 Phylogenetic Taxonomy of Living Vertebrates A phylogenetic taxonomy of living vertebrates is given below as an indented list; no fossils are included (yet). A quot;phylogeneticquot; taxonomy means that the tree can be derived exactly from this list, and vice-versa. The placement of certain fossils will be discussed as they become relevant to the course. Notice that no ranks (e.g., order, subclass) are used. The cladistic classification (p. 699) in Kardong is an attempt to do the same thing, but it is not very well done, so please ignore it. Craniata Myxinoidea Vertebrata Petromyzontia Gnathostomata Chondrichthyes Holocephali Elasmobranchii Osteichthyes Actinopterygii Brachiopterygii Unnamed taxon Chondrostei Neopterygii Lepisosteus Unnamed taxon Amia Teleostei Sarcopterygii Actinistia Choanata Dipnoi Tetrapoda Amphibia Gymnophiona Batrachia Anura Urodela Amniota Mammalia Monotremata Theria Marsupialia Eutheria Reptilia Testudines Sauria Lepidosauria Sphenodontia Squamata Archosauria Crocodylia Aves
  • 4. Comparative Anatomy page 4 Hemichordata Nonmonophyletic groups, I Urochordata Cephalochordata Myxinoidea Petromyzontia quot;Invertebrataquot; Chondrichthyes Actinopterygii Actinistia Dipnoi Amphibia Mammalia quot;Piscesquot; Testudinata quot;Crossopterygiiquot; quot;Homothermiaquot; Lepidosauria quot;Reptiliaquot; Crocodylia Aves
  • 5. Comparative Anatomy page 5 Hemichordata Nonmonophyletic groups, II Urochordata Cephalochordata Myxinoidea Petromyzontia Chondrichthyes Actinopterygii Actinistia Dipnoi quot;Agnathaquot; Amphibia Mammalia quot;Osteichthyesquot; quot;Sarcopterygiiquot; Testudinata quot;Amphibiaquot; Lepidosauria quot;Labyrinthodontiaquot; quot;Synapsidaquot; Crocodylia Aves
  • 6. Comparative Anatomy page 6 Numbers of Living Species of Chordates There are about 50,000 species of living vertebrates; this estimate varies, primarily depending on how many species of teleost fishes are recognized. Clade Name Species Common Name Hemichordata 100 acorn worms, pterobranchs Urochordata (tunicates) 2000 sea squirts, larvaceans Cephalochordata 45 amphioxus or lancelets Myxinoidea 30 hagfish Petromyzontia 40 lampreys Chondrichthyes Elasmobranchii 670 sharks, skates, rays Holocephali 30 ratfish or chimaeras Actinopterygii Brachiopterygii 13 reedfish or bichirs Chondrostei 22 sturgeon, paddlefish Lepistosteus 7 gar Amia 1 bowfin or choupique Teleostei 25000 tarpon, herring, perch, catfish, minnows, bass, etc. Actinistia 1 (maybe coelacanth (Latimeria) 2) Dipnoi 6 lungfish Amphibia Gymnophiona *160 caecilians Caudata *350 salamanders Anura 5000 frogs Mammalia Monotremata 6 platypus, echidna Marsupialia 240 opposum, kangaroos, koalas, wombats, etc. Eutheria 4300 mice, elephants, horses, whales, bats, cats, etc. Testudinata +230 turtles, tortoises, terrapins Lepidosauria Sphenodontia 2 tuatara (Sphenodon) Squamata lizards +3000 lizards Ophidia +2330 snakes Amphisbaenia +140 amphisbaenians Aves 8700 birds Crocodylia +21 alligators, crocodiles, gavials Numbers from Walker, 1987 and Wake, 1979, except as noted. *Duellman and Trueb, 1986 +Dowling and Duellman, 1978
  • 7. Comparative Anatomy page 7 Geological Time Scale If the entire geological history of earth were set to a one-year scale, the earth formed on 1 January, life appeared on 4 April, and Homo sapiens did not appear until very late on New Year's Eve. One-Year Scale Real Time Event 1 Jan 4.6 by Origin of the earth 4 Apr 3.8 by Life appears (bacteria) 9 Nov 650 my Marine invertebrates appear 20 Nov 520 my Vertebrata 29 Nov 415 my Ostracoderms 30 Nov 410 my Gnathostomata (acanthodians) Actinopterygii 2 Dec 380 my Chondrichthyes 3 Dec 360 my Tetrapoda 7 Dec 310 my Amniota 16 Dec 190 my Mammalia 20 Dec 150 my Aves 26 Dec 65 my Primates 29 Dec 30 my Anthropoidea (monkeys and apes) 11:45 pm 31 Dec 125-250,000 yr Homo sapiens Check out these web pages on the Tree of Life (ag.arizona.edu/tree/) • Vertebrates • Gnathostomata • Terrestrial vertebrates (Tetrapods) Updated by David Cannatella, September 18, 2001