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Metaves and Coronaves in a morph analysis



Thanks to Mayr for a copy of his NEXUS file, which made this much less
tedious.

The most extensive published morphological analysis of neornithines is Mayr
and Clarke (2003).  Their topology is (2 mpt's, 720 steps)-
|--Apsaravis
|--Ichthyornis
`--+--Hesperornis
   `--+--+--Tinamidae
      |  `--+--Rheidae
      |     `--Apterygidae
      `--+--+--Galliformes
         |  `--+--Anhimidae
         |     `--Anatidae
         `--+--+--Rallidae
            |  `--+--Psophiidae
            |     `--Gruidae
            `--+--Otididae
               `--+--+--Balaenicipitidae
                  |  `--+--Eurypygidae
                  |     `--Ardeidae
                  `--+--+--Threskiornithidae
                     |  `--+--Ciconiidae
                     |     `--Phoenicopteridae
                     `--+--+--Cathartidae
                        |  `--+--Sagittariidae
                        |     `--+--Strigiformes
                        |        `--+--Accipitridae
                        |           `--Falconidae
                        `--+--+--Recurvirostridae
                           |  `--Burhinidae
                           `--+--+--+--Phaethontidae
                              |  |  `--+--Fregatidae
                              |  |     `--Phalacrocoracidae
                              |  `--+--Procellariidae
                              |     `--+--Spheniscidae
                              |        `--+--Gaviidae
                              |           `--Podicepedidae
                              `--+--+--Pteroclidae
                                 |  `--Columbidae
                                 |--+--+--Opisthocomidae
                                 |  |  `--Cariamidae
                                 |  `--+--Musophagidae
                                 |     `--Cuculidae
                                 `--+--+--Aegothelidae
                                    |  `--+--Steatornithidae
                                    |     `--Trogonidae
                                    `--+--Passeriformes
                                       `--+--Coraciidae
                                          `--+--Coliidae
                                             `--Psittacidae
I ran the analysis with three ordered characters, as suggested by the
authors.  Notably, I got trees 100 steps less than reported by them, which I
assume is caused by a typo.

I wondered what would happen if I enforce Metaves and Coronaves (Fain and
Houde, 2004).  If I do, the following occurs (3 mpt's, 759 steps)-
|--Apsaravis
|--Hesperornis
`--+--Ichthyornis
   `--+--+--Tinamidae
      |  `--+--Rheidae
      |     `--Apterygidae
      `--+--+--Galliformes
         |  `--+--Anhimidae
         |     `--Anatidae
         `--+--+--+--Podicepedidae
            |  |  `--Phoenicopteridae
            |  `--+--Eurypygidae
            |     `--+--+--Opisthocomidae
            |        |  `--Phaethontidae
            |        `--+--+--Pteroclidae
            |           |  `--Columbidae
            |           `--+--Steatornithidae
            |              `--Aegothelidae
            `--+--+--+--Gaviidae
               |  |  `--Spheniscidae
               |  `--+--Procellariidae
               |     `--+--Burhinidae
               |        `--Recurvirostridae
               `--+--+--Cariamidae
                  |  |--Otididae
                  |  |--Rallidae
                  |  `--+--Psophiidae
                  |     `--Gruidae
                  `--+--Ardeidae
                     `--+--+--+--Threskiornithidae
                        |  |  `--Ciconiidae
                        |  `--+--Balaenicipitidae
                        |     `--+--Fregatidae
                        |        `--Phalacrocoracidae
                        `--+--+--Cathartidae
                           |  `--Saggitariidae
                           `--+--+--Strigiformes
                              |  `--+--Accipitridae
                              |     `--Falconidae
                              `--+--+--Psittacidae
                                 |  `--+--Coliidae
                                 |     `--Musophagidae
                                 `--+--Cuculidae
                                    `--+--Coraciidae
                                       `--+--Trogonidae
                                          `--Passeriformes
Note for this and the other enforced analyses, the non-neoavian portion was
enforced in the traditional way.  I only enforced the metavian and
coronavian clades here, not their respective subclades.

Livezey and Zusi's (2001) morphological analysis is much larger than Mayr
and Clarke's, but also unfinished and lacking a published matrix.
Constraining for their topology led to 2 mpt's of 823 steps.
|--Apsaravis
|--Hesperornis
`--+--Ichthyornis
   `--+--+--Tinamidae
      |  `--+--Rheidae
      |     `--Apterygidae
      `--+--+--Galliformes
         |  `--+--Anhimidae
         |     `--Anatidae
         `--+--Cariamidae
            `--+--+--Phaethontidae
               |  `--+--Fregatidae
               |     `--+--Balaenicipitidae
               |        `--Phalacrocoracidae
               `--+--+--+--Steatornithidae
                  |  |  `--+--Ardeidae
                  |  |     `--+--Ciconiidae
                  |  |        `--Threskiornithidae
                  |  `--+--Cathartidae
                  |     `--+--Sagittariidae
                  |        `--+--Falconidae
                  |           `--Accipitridae
                  `--+--Otididae
                     `--+--+--Burhinidae
                        |  `--+--+--+--Psophiidae
                        |     |  |  `--Opisthocomidae
                        |     |  `--Gruidae
                        |     `--+--Eurypygidae
                        |        `--+--Phoenicopteridae
                        |           `--+--Podicepedidae
                        |              `--+--Gaviidae
                        |                 `--+--Spheniscidae
                        |                    `--Procellariidae
                        `--+--+--+--Pteroclidae
                           |  |  `--Columbidae
                           |  `--Recurvirostridae
                           `--+--Musophagidae
                              `--+--Coliidae
                                 `--+--Rallidae
                                    `--+--Psittacidae
                                       `--+--Passeriformes
                                          `--+--Trogonidae
                                             `--+--Cuculidae
                                                `--+--Strigiformes
                                                   `--Aegothelidae
Note- for this constraint, I couldn't specify the placement of Coraciidae,
because Livezey and Zusi's tree has Coraciiformes as polyphyletic and
doesn't indicate which branch contains coraciids.  They end up either sister
to coliids or strigiformes+aegothelids in the constrained analysis.  Livezey
and Zusi have no coraciiformes in the former spot, but do have some in the
latter.

How about constraining traditional orders?  That makes 5 mpt's of 744 steps.
|--Apsaravis
|--Hesperornis
`--+--Ichthyornis
   `--+--+--Tinamidae
      |  `--+--Rheidae
      |     `--Apterygidae
      `--+--+--Galliformes
         |  `--+--Anhimidae
         |     `--Anatidae
         `--+--+--Opisthocomidae
            |  `--+--Cuculidae
            |     `--Musophagidae
            `--+--+--+--Burhinidae
               |  |  `--Recurvirostridae
               |  `--+--+--+--Balaenicipitidae
               |     |  |  `--Ardeidae
               |     |  `--+--Threskiornithidae
               |     |     `--+--Ciconiidae
               |     |        `--Phoenicopteridae
               |     `--+--+--Rallidae
               |        |  `--+--Cariamidae
               |        |     `--Otididae
               |        `--+--Eurypygidae
               |           `--+--Gruidae
               |              `--Psophiidae
               `--+--+--+--Phaethontidae
                  |  |  `--+--Fregatidae
                  |  |     `--Phalacrocoracidae
                  |  `--+--Procellariidae
                  |     `--+--Spheniscidae
                  |        `--+--Gaviidae
                  |           `--Podicipedidae
                  |--+--Strigiformes
                  |  |--Cathartidae
                  |  |--Sagittariidae
                  |  `--+--Accipitridae
                  |     `--Falconidae
                  |--+--Psittacidae
                  |  `--+--Pteroclidae
                  |     `--Columbidae
                  `--+--Coraciidae
                     |--+--Trogonidae
                     |  `--Passeriformes
                     `--+--Coliidae
                        `--+--Aegothelidae
                           `--Steatornithidae
This resembles Cracraft's infamous 1988 study in some ways, including...
-
(Pelecaniformes,Procellariiformes(Sphenisciformes(Gaviiformes,Podicepidiform
es)))
- (Falconiformes,Strigiformes)
- (Gruiformes,Charadriiformes,Ardeidae), though Columbiformes was in this
clade in Cracraft's study.
- (Coraciiformes,Passeriformes)
Note Opisthocomus was allowed to fall where it wanted, due to its
historically controversial position.

Sibley and Alquist's (1990) oft maligned topology is 861 steps.

So, according to parsimony, the classic topology is only 24 more steps than
Mayr and Clarke's tree, though it is thought to be incorrect by most
ornithologists.  Fain and Houde's new hypothesis is only 39 steps longer,
despite appearing to be starkingly novel.  Livezey and Zusi's topology,
which seems like it should be the best morphology-based one, is a whopping
103 more steps.  Interesting how our intuitions about how probable various
topologies are sometimes aren't reflected by parsimony.

Mickey Mortimer
Undergraduate, Earth and Space Sciences
University of Washington
The Theropod Database - http://students.washington.edu/eoraptor/Home.html