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Applying Sereno's definitions to Neotetanurae Part 1
Because I've already extensively researched neotetanurine phylogenies for my
studies, I figured they were a good group to test Taxon Search's active
definitions on. Below, I list all published neotetanurine phylogenies based
on available matrices (plus a few from theses), and apply Taxon Search's
active definitions to them. This will demonstrate where compositional or
taxonomic conflict exists.
*please let the formatting survive, so that the tens of cladograms are
readable...*
Gauthier, 1986
Neotetanurae
Tyrannoraptora
|--Allosauroidea
| Tyrannosauroidea
| Allosauridae
| |--Allosauridae
| `--Tyrannosauridae
`--Coelurosauria
Maniraptoriformes
|--Ornithomimiformes
| Ornithomimosauria
`--Maniraptora
|--Coelurus
|--Ornitholestes
|--Compsognathidae
| Compsognathus
|--Caenagnathidae
| Elmisauridae
|--Oviraptoriformes
| Oviraptorosauria
| Oviraptoridae
| Caenagnathoidea sensu stricto
|--Saurornitholestes
`--Paraves
Eumaniraptora
|--Aves
`--Deinonychosauria
|--Dromaeosauridae
`--Troodontidae
This shows Tyrannosauroidea needs Allosaurus fragilis as an external
specifier, and Allosauridae
needs Tyrannosaurus rex as an external specifier. Also, it indicates a
problem with using
Chirostenotes (an elmisaurid in Gauthier's usage) as an internal specifier
for Caenagnathidae,
when we have the topology (Chirostenotes (Caenagnathus, Oviraptor)). Does
Neotetanurae or
Tyrannoraptora get priority if a (Passer (Sinraptor, Carcharodontosaurus,
Allosaurus,
Tyrannosaurus)) topology exists? Both are Sereno's names, but I say
Neotetanurae since it was
named earlier.
Chatterjee, 1991
|--other theropods
`--Aves
|--Archaeopterygidae
| Archaeopteryx
`--Ornithurae
|--Avimimus
`--+--Protoavis
`--Ornithothoraces
|--Enantiornithes
| Gobipteryx (+Sinornis?)
|--Hesperornithes
| Hesperornis
`--Euornithes
Ichthyornis (+Passer)
Holtz, 1992
Neotetanurae
|--Allosauroidea
| |--Megalosaurus
| |--Piatnitzkysaurus
| `--+--Eustreptospondylinae
| | Eustreptospondylus
| `--+--Allosauridae
| | Allosaurus
| |--Carcharodontosauridae
| | Acrocanthosaurus (+Carcharodontosaurus?)
| `--Chilantaisaurus
`--Coelurosauria
|--Compsognathidae
| Compsognathus
`--+--Ornitholestes
`--Tyrannoraptora
Maniraptoriformes
|--Maniraptora
| Paraves
| Eumaniraptora
| |--Dromaeosauridae
| `--Aves
| Archaeopteryx (+Passer?)
`--Ornithomimiformes
Oviraptoriformes
Caenagnathoidea
|--Oviraptorosauria
| Oviraptoridae
`--Caenagnathidae
|--Elmisauridae
`--+--Troodontidae
`--+--Tyrannosauroidea
| Tyrannosauridae
`--Ornithomimosauria
|--Avimimus
`--+--Ornithomimidae
`--+--Harpymimus
`--Garudimimus
If we have a topology like this (Passer (Tyrannosaurus, Ornithomimus)), does
Tyrannoraptora or
Maniraptoriformes get priority? I say Maniraptoriformes because it was
named first. And if we
have (Passer (Ornithomimus, Oviraptor), does Maniraptora or Paraves get
priority? I say
Maniraptora, as it was named first. And how about Ornithomimiformes vs.
Oviraptoriformes? Both
are Sereno's names, published in the same paper, he should decide. This also
shows Caenagnathidae
needs an additional external specifier, such as Ornithomimus velox.
Sanz and Buscalioni, 1992
This excludes Enantiornithes and Ambiortus, which cause a massive
ornithurine polytomy.
|--other theropods
`--Aves
|--Archaeopterygidae
| Archaeopteryx
`--Ornithurae
|--Iberomesornis
`--Ornithothoraces
|--Concornis (+Sinornis?)
`--Euornithes
|--Hesperornithes
`--Carinatae
Here, the placement of Ornithothoraces is uncertain, because Sereno uses
Sinornis instead of
Iberomesornis.
Sanz and Bonaparte, 1992
|--other theropods
`--Aves
|--Archaeopterygidae
| Archaeopteryx
`--Ornithurae
|--Iberomesornis
`--Euornithes
Perez-Moreno et al., 1993
|--Ceratosauria
`--Tetanurae
|--Eustreptospondylinae
| Eustreptospondylus
`--+--Piatnitzkysaurus
`--Neotetanurae
|--Allosauroidea
| Allosauridae
| |--Allosaurus
| `--Monmirat theropod
`--Coelurosauria
Maniraptoriformes
Tyrannoraptora
|--Maniraptora
| Eumaniraptora
| Deinonychus (+Passer?)
`--Ornithomimiformes
|--Tyrannosauroidea
| Tyrannosauridae
| Albertosaurus (+Tyrannosaurus)
`--Ornithomimosauria
Ornithomimidae
Gallimimus (+Ornithomimus)
Tyrannoraptora vs. Maniraptoriformes again.
Chiappe and Calvo, 1994 and modifications
This is the data from the original paper, plus Norell et al. (1996) and
Zhang and Zhou (2000)
combined.
Eumaniraptora
|--Dromaeosauridae
| Velociraptorinae
`--Aves
|--Archaeopterygidae
| Archaeopteryx
`--Ornithurae
|--Iberomesornis
`--+--Alvarezsaurus
|--Alvarezsauridae
| |--Patagonykus
| `--Mononykinae
| Mononykus (+Shuvuuia)
`--+--Confuciusornithidae
| Confuciusornis
`--+--Protopteryx
`--Ornithothoraces
|--Enantiornithes
| Sinornis
`--Euornithes
|--+--Concornis
| `--Enantiornis
`--+--Patagopteryx
`--+--Chaoyangia
`--+--Hesperornithes
`--Carinatae
|--Ichthyornis
`--Neornithes
Using Shuvuuia as the internal specifier for Alvarezsauridae leaves
Alvarezsaurus potentially out
of the clade. Using Sinornis as the internal specifier for Enantiornithes
leaves Enantiornis out
of the clade. Here, the use of Sinornis as a specifier for Ornithothoraces
is actually
beneficial.
Chiappe and Calvo, 1994 and Norell et al., 1996
These have compatable topologies.
Eumaniraptora
|--Dromaeosauridae
| Velociraptorinae
`--Aves
|--Archaeopterygidae
| Archaeopteryx
`--Ornithurae
|--Alvarezsauridae
| |--Alvarezsaurus
| `--+--Patagonykus
| `--Mononykinae
| Mononykus (+Shuvuuia)
`--+--Iberomesornis
`--Ornithothoraces
|--Enantiornithes
| |--Concornis
| `--Enantiornithes sensu stricto (+Sinornis)
`--Euornithes
|--Patagopteryx
`--+--Hesperornithes
`--Carinatae
|--Ichthyornis
`--Neornithes
Here Ornithothoraces doesn't apply to the same node Chiappe intended when he
named it, as Sereno uses Sinornis as an internal specifier instead of
Iberomesornis.
Zhang and Zhou, 2000
Eumaniraptora
|--Dromaeosauridae
`--Aves
|--Archaeopterygidae
| Archaeopteryx
`--Ornithurae
|--Confuciusornithidae
| Confuciusornis
`--Ornithothoraces
|--Enantiornithes
| |--Protopteryx
| `--+--Sinornis
| `--+--Concornis
| `--Enantiornis
`--Euornithes
|--Chaoyangia
`--+--Hesperornithes
`--Carinatae
|--Ichthyornis
`--Neornithes
Lu et al., 2002
Eumaniraptora
|--Dromaeosauridae
| Velociraptorinae
`--Aves
|--Archaeopterygidae
| Archaeopteryx
`--Ornithurae
|--Caudipteryx
`--+--Alvarezsaurus
`--+--Alvarezsauridae
| |--Patagonykus
| `--Mononykinae
| Mononykus (+Shuvuuia)
`--+--Oviraptoriformes
| Oviraptorosauria
| Oviraptoridae
`--Paraves
|--Iberomesornis
`--+--Confuciusornithidae
| Confuciusornis
`--Ornithothoraces
|--Enantiornithes (+Sinornis)
`--Euornithes
|--Patagopteryx
`--+--Hesperornithes
`--Carinatae
|--Ichthyornis
`--Neornithes
Using Shuvuuia as the internal specifier for Alvarezsauridae leaves
Alvarezsaurus out of the
clade.
Holtz, 1994
Neotetanurae
|--Allosauroidea
| |--Carcharodontosauridae
| | Acrocanthosaurus (+Carcharodontosaurus?)
| `--Allosauridae
| Allosaurus
`--Coelurosauria
|--Compsognathidae
| Compsognathus
`--+--Ornitholestes
`--Tyrannoraptora
Maniraptoriformes
|--Maniraptora
| Paraves
| Eumaniraptora
| |--Dromaeosauridae
| `--Aves
| Archaeopteryx (+Passer)
`--Ornithomimiformes
Oviraptoriformes
Caenagnathoidea
|--Oviraptorosauria
| Oviraptoridae
`--Caenagnathidae
|--Elmisauridae
`--+--Avimimus
`--+--Tyrannosauroidea
| Tyrannosauridae
`--+--Ornithomimosauria
`--Troodontidae
The same comments apply as for Holtz, 1992.
Perez-Moreno et al., 1994
Neotetanurae
|--Allosauroidea
| Allosauridae
| Allosaurus
`--Coelurosauria
Tyrannoraptora
|--Tyrannosauroidea
| Tyrannosauridae
| Albertosaurus (+Tyrannosaurus)
`--Maniraptoriformes
|--Maniraptora
| Eumaniraptora
| Deinonychus (+Passer?)
`--Ornithomimiformes
|--Troodontidae
`--Ornithomimosauria
|--Pelecanimimus
`--+--Garudimimus
`--Ornithomimidae
Gallimimus (+Ornithomimus)
Russell and Dong, 1994
This incorporates two possible positions for Passer, as no Aves were
analyzed.
|--Coelophysidae
`--Maniraptoriformes
|--Maniraptora
| Paraves
| |--Spinosauridae
| `--Neotetanurae
| |--Sinraptoridae
| `--+--Allosauroidea
| | Allosauridae
| | Allosaurus
| `--Coelurosauria
| Tyrannoraptora
| |--Tyrannosauroidea
| | Tyrannosauridae
| `--Eumaniraptora
| Dromaeosauridae (+Passer?)
`--Ornithomimiformes
Oviraptoriformes
|--Ornithomimosauria
`--Troodontidae
|--Therizinosauria
`--+--Oviraptorosauria
`--Troodontidae sensu stricto
Troodontidae needs Oviraptor as an external specifier.
|--Coelophysidae
`--Neotetanurae
Tyrannoraptora
Eumaniraptora
|--Allosauroidea
| Dromaeosauridae
| |--Spinosauridae
| `--+--Sinraptoridae
| `--Allosauridae
| |--Allosaurus
| `--+--Tyrannosauroidea
| | Tyrannosauridae
| `--Dromaeosauridae sensu stricto
`--Coelurosauria
Maniraptoriformes
|--Ornithomimiformes
| Ornithomimosauria
`--Maniraptora
|--Therizinosauria
`--+--Oviraptoriformes
| Oviraptorosauria
`--Paraves
Troodontidae (+Passer?)
Again we have Neotetanurae vs. Tyrannoraptora, but this time Eumaniraptora
joins in. This can be
solved here by having Eumaniraptora by definition exclude Ornithomimus,
which forces it to be a
subset of Maniraptora if it exists. Dromaeosauridae needs Tyrannosaurus as
an external specifier,
and Allosauridae again needs Tyrannosaurus as one.
Makovicky, 1995
Neotetanurae
|--Allosauroidea
| Allosauridae
| Allosaurus
`--Coelurosauria
|--Coelurus
`--Tyrannoraptora
|--Tyrannosauroidea
| Tyrannosauridae
`--Maniraptorifrmes
|--Maniraptora
| Paraves
| |--Ornitholestes
| `--Eumaniraptora
| |--Dromaeosauridae
| `--Aves
`--Ornithomimiformes
Oviraptoriformes
|--+--Therizinosauria
| `--Oviraptorosauria
`--+--Troodontidae
`--Ornithomimosauria
|--Ornithomimidae
`--Avimimus
Maniraptora vs. Paraves, and Ornithomimiformes vs. Oviraptoriformes again.
Hou et al., 1996
|--other theropods
`--Aves
Ornithothoraces
|--Archaeopterygidae
| Confuciusornithidae
| Enantiornithes
| |--Archaeopteryx
| `--+--Confuciusornis
| `--Sinornis
`--Ornithurae
Euornithes
|--Liaoningornis
`--+--Chaoyangia
`--+--Hesperornithes
| Hesperornis
`--Carinatae
Ichthyornis (+Passer)
Here we see that Archaeopterygidae needs Confuciusornis as an external
specifier,
Confuciusornithidae needs Sinornis as one, and Enantiornithes Confuciusornis
as one. Also, Aves
and Ornithorthoraces are synonymous, as are Ornithurae and Euornithes. I
recommend keeping Aves and Ornithurae based on historical priority and
amount of use.
Novas, 1996
Tyrannoraptora
Maniraptoriformes
|--Tyrannosauroidea
| Tyrannosauridae
|--Ornithomimiformes
| Ornithomimosauria
| Ornithomimidae
`--Maniraptora
Eumaniraptora
|--Dromaeosauridae
| Deinonychus (+Dromaeosaurus?)
`--Aves
|--Archaeopterygidae
| Archaeopteryx
`--Ornithurae
|--Alvarezsauridae
| |--Alvarezsaurus
| `--+--Patagonykus
| `--Mononykinae
| Mononykus (+Shuvuuia)
`--other ornithurines
Sereno et al., 1996
Neotetanurae
|--Allosauroidea
| |--Sinraptoridae
| |--Cryolophosaurus
| |--Monolophosaurus
| |--Allosauridae
| | Allosaurus
| `--Carcharodontosauridae
| |--Giganotosaurus
| `--+--Acrocanthosaurus
| `--Carcharodontosaurus
`--Coelurosauria
|--Ornitholestes
`--+--Deltadromeus
`--other coelurosaurs
Martin, 1997
Tyrannoraptora
Maniraptoriformes
Eumaniraptora
|--Paraves
| Maniraptora
| Aves
| |--Archaeopterygidae
| | Archaeopteryx
| `--Ornithurae
| Carinatae
| Ichthyornis
`--Ornithomimiformes
Oviraptoriformes
|--all zero outgroup
`--+--Dromaeosauridae
| |--Tyrannosauroidea
| | Tyrannosaurus
| `--Deinonychus (+Dromaeosaurus?)
`--+--Troodontidae
| Troodon
`--+--Oviraptorosauria
| Oviraptor
`--+--Alvarezsauridae
| Mononykinae
| Mononykus (+Shuvuuia)
|--Ornithomimosauria
| Ornithomimus
`--Struthiomimus
It's Tyrannoraptora vs. Maniraptoriformes vs. Eumaniraptora again. This
time we also have
Maniraptora vs. Paraves, and Ornithomimiformes vs. Oviraptoriformes. This
again shows
Dromaeosauridae needs Tyrannosaurus as an external specifier.
Sues, 1997
Neotetanurae
|--Allosauroidea
| Allosauridae
| Allosaurus
`--Coelurosauria
Tyrannoraptora
|--Tyrannosauroidea
| Tyrannosauridae
`--Maniraptoriformes
|--Ornithomimiformes
| Ornithomimosauria
| Ornithomimidae
`--Maniraptora
|--Oviraptoriformes
| |--Therizinosauria
| | Therizinosauroidea
| `--Oviraptorosauria
| Caenagnathoidea
`--Paraves
Deinonychosauria
|--Troodontidae
`--Dromaeosauridae (+Passer?)
Chiappe et al., 1998 and Ji et al., 1998
Eumaniraptora
|--Protarchaeopteryx
|--Dromaeosauridae
| Velociraptorinae (+Dromaeosaurus?)
`--+--Caudipteryx
`--Aves
|--Archaeopterygidae
| Archaeopteryx
`--Ornithurae
|--Alvarezsauridae
| |--Alvarezsaurus
| `--+--Patagonykus
| `--Mononykinae
| |--Mononykus
| |--Shuvuuia
| `--Parvicursor
`--Ornithothoraces
|--Enantiornithes (+Sinornis?)
`--Euornithes
Forster et al., 1998 and Zhang et al., 2002
This is the result when Epidendrosaurus is added to Forster et al.'s
complete matrix, and thus
differs from Zhang et al.'s tree, which excluded Unenlagia.
Neotetanurae
|--Allosauroidea
| Allosauridae
| Allosaurus
`--Coelurosauria
|--Compsognathidae
| Compsognathus
`--Maniraptoriformes
|--Ornithomimiformes
| Ornithomimosauria
| Ornithomimidae
`--Maniraptora
|--Oviraptoriformes
| Oviraptorosauria
| Oviraptoridae
`--Paraves
Eumaniraptora
Tyrannoraptora
|--Dromaeosauridae
| |--Tyrannosauroidea
| | Tyrannosauridae
| `--Dromaeosauridae sensu stricto
`--+--Troodontidae
`--+--Epidendrosaurus
`--Aves
|--Archaeopterygidae
| Unenlagiinae
| |--Unenlagia
| `--+--Archaeopteryx
| `--Rahonavis
`--Ornithurae
|--Alvarezsauridae
`--+--Iberomesornis
`--Ornithothoraces
|--Enantiornithes (+Sinornis)
`--Euornithes
|--Patagopteryx
`--other euornithines
Eumaniraptora vs. Tyrannoraptora again. Dromaeosauridae needing to exclude
Tyrannosaurus too.
Unenlagiinae needs to exclude Archaeopteryx, though it can be a subfamily of
Archaeopterygidae.
Again, Ornithothoraces doesn't define the clade Chiappe intended if you use
Sinornis as a
specifier.
Makovicky and Sues, 1998
Neotetanurae
|--Allosauroidea
| Allosauridae
| Allosaurus
`--Coelurosauria
Tyrannoraptora
|--Tyrannosauroidea
| Tyrannosauridae
`--Maniraptoriformes
|--Ornithomimiformes
| Ornithomimosauria
`--Maniraptora
|--Ornitholestes
`--+--Coelurus
`--+--Paraves
| Eumaniraptora
| |--Aves
| `--Deinonychosauria
| |--Dromaeosauridae
| `--Troodontidae
`--Oviraptoriformes
|--Therizinosauria
| Therizinosauroidea
`--Oviraptorosauria
Caenagnathoidea
|--Caenagnathidae
|--Microvenator
`--Oviraptoridae
Chatterjee, 1999
Eumaniraptora
|--Dromaeosauridae
`--Aves
|--Archaeopterygidae
| Archaeopteryx
`--Ornithurae
|--Alvarezsauridae
| |--Mononykinae
| | Mononykus (+Shuvuuia?)
| `--Avimimus
`--Ornithothoraces
|--Enantiornithes
| Sinornis
`--Euornithes
|--Iberomesornis
`--+--Protoavis
`--+--"Cathayornis"
`--+--Enantiornithes sensu stricto
`--+--Hesperornithes
| |--Patagopteryx
| `--Hesperornithes sensu stricto
`--Carinatae
|--+--Ambiortus
| `--Ichthyornithes
`--Neornithes
This again shows that Enantiornithes needs Enantiornis as the internal
specifier, not Sinornis.
And it's another case where Ornithothoraces doesn't define the clade Chiappe
intended if you use
Sinornis as a specifier.
Sereno, 1999
This combines the results from Sereno's Tetanurae and Aves analyses.
Neotetanurae
|--Allosauroidea
| |--Sinraptoridae
| `--Allosauridae
`--Coelurosauria
|--Deltadromeus
|--Ornitholestes
|--Compsognathidae
`--Maniraptoriformes
|--Ornithomimiformes
| |--Therizinosauria
| `--+--Alvarezsauridae
| `--Ornithomimosauria
`--Maniraptora
Tyrannoraptora
|--Tyrannosauroidea
`--+--Oviraptoriformes
| Oviraptorosauria
| |--Caudipteryx
| `--Caenagnathoidea
| |--Caenagnathidae
| `--Oviraptoridae
`--Paraves
Eumaniraptora
|--Deinonychosauria
| |--Dromaeosauridae
| `--Troodontidae
`--Aves
|--Archaeopterygidae
| Archaeopteryx
`--Ornithurae
|--Confuciusornithidae
| Confuciusornis
`--Ornithothoraces
|--Enantiornithes (+Sinornis)
`--Euornithes
In a case like this, where neither Tyrannosaurus, Ornithomimus,
Therizinosaurus nor Troodon are
closer to Oviraptor than Passer is, Oviraptorosauria should have priority
over Oviraptoriformes.
Xu et al., 1999
Maniraptoriformes
|--Ornithomimiformes
| Ornithomimosauria
| Ornithomimidae
`--Maniraptora
|--Paraves
| Dromaeosauridae (+Passer?)
`--Oviraptoriformes
Troodontidae
|--Troodontidae sensu stricto
`--+--Oviraptorosauria
`--Therizinosauria
Therizinosauroidea
|--Beipiaosaurus
`--+--Alxasaurus
`--Therizinosauridae
Again, Troodontidae needs Oviraptor as an external specifier.
Xu et al., 1999
Neotetanurae
|--Allosauroidea
| Allosauridae
| Allosaurus
`--Coelurosauria
|--Compsognathidae
| Compsognathus
`--Maniraptoriformes
|--Ornithomimiformes
| Ornithomimosauria
| Ornithomimidae
`--Maniraptora
|--Oviraptoriformes
| Oviraptorosauria
| Oviraptoridae
`--Paraves
|--Caudipteryx
`--+--Protarchaeopteryx
`--+--Troodontidae
`--Eumaniraptora
|--Aves
| |--Archaeopterygidae
| | Archaeopteryx
| `--Ornithurae
| Rahonavis (+Passer?)
`--Dromaeosauridae
|--Microraptorinae
| Microraptor
`--+--Sinornithosaurus
|--Dromaeosaurinae
| Dromaeosaurus
`--Velociraptorinae
Zhou, 1999
Eumaniraptora
|--Deinonychosauria
`--Aves
|--Archaeopteryx
`--Ornithurae
|--Confuciusornis
`--Ornithothoraces
|--Enantiornithes
| |--Eocathayornis
| |--Sinornis
| `--Enantiornis
`--Euornithes
|--Liaoningornis
`--+--Chaoyangia
`--+--Gansus
`--Carinatae
|--Ichthyornis
`--Neornithes
Azuma and Currie, 2000
Neotetanurae
|--Allosauroidea
| |--Fukuiraptor
| `--+--Sinraptoridae
| | Sinraptor
| `--+--Allosauridae
| | Allosaurus
| `--Carcharodontosauridae
| Acrocanthosaurus (+Carcharodontosaurus?)
`--Coelurosauria
|--Compsognathidae
| Sinosauropteryx (+Compsognathus?)
`--Maniraptoriformes
Tyrannoraptora
|--Ornithomimiformes
| |--Tyrannosauroidea
| | Tyrannosauridae
| `--Ornithomimosauria
| Ornithomimidae
`--Maniraptora
|--Oviraptoriformes
| Oviraptorosauria
| Oviraptoridae
`--Paraves
Eumaniraptora
Dromaeosauridae (+Passer?)
Again, we have Maniraptoriformes vs. Tyrannoraptora.
Currie and Carpenter, 2000
Neotetanurae
|--Allosauroidea
| |--Monolophosaurus
| `--+--Sinraptoridae
| |--Allosauridae
| | |--Allosaurus
| | `--Acrocanthosaurus
| `--Carcharodontosauridae
| |--Giganotosaurus
| `--Carcharodontosaurus
`--Coelurosauria
Tyrannoraptora
|--Tyrannosauroidea
| Tyrannosauridae
`--+--Oviraptoriformes
| Oviraptorosauria
`--Paraves
Eumaniraptora
Dromaeosauridae (+Passer?)
Holtz, 2000 and 2001
This was taken from the matrix of his 2001 analysis, and differs in
unpublished ways from the
2000 topology.
Neotetanurae
|--Allosauroidea
| |--Sinraptoridae
| | |--Sinraptor
| | `--Yangchuanosaurus
| `--+--Allosauridae
| | |--Allosaurus
| | `--Neovenator
| `--Carcharodontosauridae
| |--Acrocanthosaurus
| `--+--Carcharodontosaurus
| `--Giganotosaurus
`--Coelurosauria
|--Gasosaurus
|--Proceratosaurus
`--+--+--Bagaraatan
| `--Ornitholestes
`--+--Scipionyx
`--Maniraptoriformes
Tyrannoraptora
|--Ornithomimiformes
| Compsognathidae
| |--Compsognathidae sensu lato
| `--+--Tyrannosauroidea
| | Tyrannosauridae
| `--+--Troodontidae
| `--Ornithomimosauria
| |--Pelecanimimus
| `--Ornithomimidae
`--Maniraptora
|--Dryptosaurus
`--+--Coelurus
`--+--Deltadromeus
|--Protarchaeopteryx
|--Caudipteryx
`--+--Oviraptoriformes
| |--Therizinosauria
| `--Oviraptorosauria
| Caenagnathoidea
| |--Caenagnathidae
| |--Microvenator
| `--Oviraptoridae
`--Paraves
Eumaniraptora
|--Dromaeosauridae
`--Aves
|--Archaeopterygidae
| Archaeopteryx
|--Unenlagiinae
| |--Unenlagia
| `--Rahonavis
`--Ornithurae
|--Alvarezsauridae
`--+--Confuciusornithidae
| Confuciusornis
`--Ornithothoraces
Holtz, 2000
This had two most parsimonious topologies (when restricted to the taxa
below), differing in the
position of Troodontidae.
Neotetanurae
|--Allosauroidea
| |--Monolophosaurus
| `--+--Sinraptoridae
| | |--Sinraptor
| | `--Yangchuanosaurus
| `--+--Allosauridae
| | |--Allosaurus
| | `--Neovenator
| `--Carcharodontosauridae
| |--Acrocanthosaurus
| `--+--Carcharodontosaurus
| `--Giganotosaurus
`--Coelurosauria
|--Gasosaurus
`--+--Dryptosaurus
`--+--Ornitholestes
`--+--Coelurus
`--+--Scipionyx
`--+--Bagaraatan
`--Maniraptoriformes
Tyrannoraptora
|--Ornithomimiformes
| |--Tyrannosauroidea
| | Tyrannosauridae
| `--+--Troodontidae
| `--Ornithomimosauria
| |--Pelecanimimus
| `--Ornithomimidae
`--Maniraptora
|--Compsognathidae
`--+--Oviraptoriformes
| |--Therizinosauria
| `--Oviraptorosauria
| Caenagnathoidea
| |--Caenagnathidae
| |--Microvenator
| `--Oviraptoridae
`--Paraves
Eumaniraptora
|--Dromaeosauridae
`--+--Rahonavis
`--Aves
|--Archaeopterygidae
| Archaeopteryx
`--Ornithurae
|--Alvarezsauridae
`--Ornithothoraces
...continued from non-maniraptoriform tree above
`--Maniraptoriformes
Tyrannoraptora
|--Ornithomimiformes
| |--Tyrannosauroidea
| | Tyrannosauridae
| `--Ornithomimosauria
| |--Pelecanimimus
| `--Ornithomimidae
`--Maniraptora
|--Compsognathidae
`--+--Oviraptoriformes
| |--Therizinosauria
| `--Oviraptorosauria
| Caenagnathoidea
| |--Caenagnathidae
| |--Microvenator
| `--Oviraptoridae
`--Paraves
|--Troodontidae
`--Eumaniraptora
|--Dromaeosauridae
`--+--Rahonavis
`--Aves
|--Archaeopterygidae
| Archaeopteryx
`--Ornithurae
|--Alvarezsauridae
`--Ornithothoraces
Tyrannoraptora vs. Maniraptoriformes again. Compsognathidae needs
Tyrannosaurus as an external
specifier.
Norell et al., 2000
Maniraptoriformes
|--Ornithomimiformes
| Ornithomimosauria
| Ornithomimidae
`--Maniraptora
|--Eumaniraptora
| |--Dromaeosauridae
| `--+--Oviraptoriformes
| | Oviraptorosauria
| `--Paraves
| Aves
`--Troodontidae
|--IGM 100/44
`--+--Sinornithoides
`--+--Byronosaurus
`--+--Saurornithoides mongoliensis
|--Saurornithoides junior
`--Troodon
Rauhut, 2000 and 2003
Neotetanurae
|--Allosauroidea
| |--Eustreptospondylinae
| | Magnosaurus (+Eustreptospondylus)
| |--Poekilopleuron
| |--Spinosauroidea
| | |--Torvosauridae
| | | Torvosaurus
| | `--Spinosauridae
| | |--Chilantaisaurus
| | `--Spinosauridae sensu stricto
| |--Monolophosaurus
| `--+--Afrovenator
| `--+--Allosauridae
| | Allosaurus
| `--+--Sinraptoridae
| `--Carcharodontosauridae
| |--Neovenator
| `--other carcharodontosaurids
`--Coelurosauria
|--Proceratosaurus
`--+--Compsognathidae
| |--Coelurus
| `--+--Sinosauropteryx
| `--+--Compsognathus
| `--Mirischia
`--Tyrannoraptora
|--Tyrannosauroidea
| |--Stokesosaurus
| `--Tyrannosauridae
`--+--Ornitholestes
`--Maniraptoriformes
|--Ornithomimiformes
| |--Deltadromeus
| `--Ornithomimosauria
`--Maniraptora
|--Bagaraatan
|--Oviraptoriformes
| |--Therizinosauria
| `--Oviraptorosauria
| |--Avimimus
| |--Microvenator
| `--Caenagnathoidea
`--Paraves
Eumaniraptora
|--Deinonychosauria
| |--Troodontidae
| `--Dromaeosauridae
`--+--Unenlagiinae
| Unenlagia
`--Aves
Chiappe, 2001 and 2002
Neotetanurae
|--Allosauroidea
`--Coelurosauria
Eumaniraptora
|--Dromaeosauridae
| Velociraptorinae (+Dromaeosaurus?)
`--+--Troodontidae
`--+--Alvarezsauridae
| |--Alvarezsaurus
| `--+--Patagonykus
| `--Mononykinae
| |--Mononykus
| `--Shuvuuia
`--Aves
|--Archaeopterygidae
| Archaeopteryx
`--Ornithurae
|--Rahonavis
`--+--Confuciusornithidae
| |--Confuciusornis
| `--Changchengornis
`--Ornithothoraces
|--Enantiornithes
| |--+--Iberomesornis
| | `--Noguerornis
| `--+--Eoalulavis
| `--+--Sinornis
| |--Concornis
| |--Neuquenornis
| `--Gobipteryx
`--Euornithes
|--Vorona
|--Patagopteryx
`--+--Hesperornithes
| Hesperornis
`--+--Ambiortus
`--Carinatae
|--Ichthyornis
`--Neornithes
Anas (+Passer)
Clarke and Chiappe, 2001
|--Confuciusornithidae
| Confuciusornis
`--Ornithothoraces
|--Enantiornithes (+Sinornis)
`--Euornithes
Carinatae
|--Ichthyornis
`--+--Limenavis
`--+--Lithornis
`--Neornithes
Holtz, 2001
Maniraptoriformes
Tyrannoraptora
|--Ornithomimiformes
| Ornithomimosauria
|--Maniraptora (+Passer?)
`--Tyrannosauroidea
|--Siamotyrannus
`--+--+--Stygivenator
| |--juvenile Daspletosaurus
| `--Alectrosaurus
`--+--Alioramus
`--+--Shanshanosaurus
`--Tyrannosauridae
|--Albertosaurinae
| |--Gorgosaurus
| `--Albertosaurus
`--Tyrannosaurinae
|--Daspletosaurus torosus
|--Daspletosaurus sp. nov.
`--+--Tarbosaurus
`--Tyrannosaurus
Norell and Clarke, 2001 and modifications
This was taken from Clarke (2004), with taxa added after Clarke (2002), Ji
et al. (2005) and Zhou
and Zhang (2005).
|--Huaxiagnathus
`--Eumaniraptora
|--Compsognathidae
| Sinosauropteryx (+Compsognathus?)
|--Protarchaeopteryx
|--Oviraptoriformes
| Caudipteryx (+Oviraptor?)
|--Troodontidae
| Sinovenator (+Troodon?)
|--Dromaeosauridae
| Sinornithosaurus (+Dromaeosaurus?,+Deinonychus?,+Velociraptor?)
|--Rahonavis
`--Paraves
|--Microraptorinae
| Microraptor
`--Aves
|--Archaeopterygidae
| |--Archaeopteryx
| `--Jinfengopteryx
`--Ornithurae
|--Shenzhouraptor
`--+--Sapeornis
`--Ornithothoraces
|--Enantiornithes
| Confuciusornithidae
| |--Confuciusornis
| `--+--Sinornis
| |--Concornis
| |--Neuquenornis
| |--Gobipteryx
| `--+--Vorona
| `--Liaoningornis sensu Clarke
`--Euornithes
|--Patagopteryx
`--+--Hongshanornis
`--+--"Archaeorhynchus"
`--+--Yanornis
`--+--Yixianornis
`--+--Apsaravis
`--+--Songlingornis
`--+--Hesperornithes
| |--Baptornis
| `--Hesperornithidae
| Hesperornis
`--Carinatae
|--Ichthyornis
`--+--Limenavis
`--+--Iaceornis
`--Neornithes
This again shows Confuciusornithidae needs Enantiornis as an external
specifier, and
Enantiornithes needs Confuciusornis as one.
Clarke, 2002, Clarke and Norell, 2002, Zhou and Zhang, 2002, Clarke, 2004
Eumaniraptora
|--Dromaeosauridae
`--Aves
|--Archaeopterygidae
| Archaeopteryx
`--Ornithurae
|--Shenzhouraptor
`--+--Sapeornis
`--+--Confuciusornithidae
| Confuciusornis
`--Ornithothoraces
|--Enantiornithes
| |--Sinornis
| |--Concornis
| |--Neuquenornis
| `--Gobipteryx
`--Euornithes
|--Vorona
`--+--Patagopteryx
`--+--Apsaravis
`--+--Hesperornithes
| |--Baptornis
| `--Hesperornithidae
| Hesperornis
`--Carinatae
|--Ichthyornis
`--+--Limenavis
|--Gansus
`--+--Iaceornis
`--Neornithes
Norell and Clarke, 2001
... continued from non-euornithine tree above
`--Euornithes
|--Patagopteryx
`--+--Hesperornithes
| |--Baptornis
| `--Hesperornithidae
| Hesperornis
`--+--Apsaravis
`--Carinatae
|--Ichthyornis
`--Neornithes
Norell et al., 2001 and modifications.
This tree is taken from Makovicky et al. (2005), incorporating the new
characters and three
therizinosaurs from Kirkland et al. (2005) and the modifications to
Archaeopteryx and Rahonavis
suggested by Mayr et al. (2005). Archaeornithomimus was excluded, as it had
an uncertain position
at the base of Maniraptoriformes.
Neotetanurae
|--Sinraptoridae
| Sinraptor
|--Allosauroidea
| Allosauridae
| Allosaurus
`--Coelurosauria
Tyrannoraptora
|--Tyrannosauroidea
| Tyrannosauridae
| |--Gorgosaurus
| `--Tyrannosaurinae
| Tyrannosaurus
`--+--Compsognathidae
| |--Compsognathus
| |--Huaxiagnathus
| `--Sinosauropteryx
`--Maniraptoriformes
|--Ornithomimiformes
| Ornithomimosauria
| |--Pelecanimimus
| `--+--Shenzhousaurus
| `--+--Harpymimus
| `--+--Garudimimus
| `--Ornithomimidae
| |--Gallimimus
| |--Struthiomimus
| `--+--Anserimimus
| `--Ornithomimus
`--Maniraptora
|--Ornitholestes
`--+--Alvarezsauridae
| |--Alvarezsaurus
| `--+--Patagonykus
| `--Mononykinae
| |--Shuvuuia
| `--Mononykus
`--+--Oviraptoriformes
| |--Therizinosauria
| | Therizinosauroidea
| | |--Erlikosaurus (+Therizinosaurus?)
| | |--Segnosaurus
| | |--Alxasaurus
| | `--+--Nothronychus
| | |--Beipiaosaurus
| | `--Falcarius
| `--Oviraptorosauria
| |--Incisivosaurus
| `--Caenagnathoidea
| |--Caudipteryx
| |--Avimimus
| |--Caenagnathidae
| | Chirostenotes
| |--Microvenator
| `--Oviraptoridae
| Oviraptorinae
| Ingeniinae
| |--Oviraptor
| |--Rinchenia
| |--Citipati
| |--Conchoraptor
| `--Ingenia
`--Paraves
Eumaniraptora
|--Aves
| |--Archaeopterygidae
| | Archaeopteryx
| `--Ornithurae
| Confuciusornithidae
| Confuciusornis (+Passer?)
`--Deinonychosauria
|--Troodontidae
| |--+--Mei
| | `--Sinovenator
| `--+--IGM 100/44
| `--+--Sinornithoides
| |--Byronosaurus
| `--+--Troodon
| `--+--Saurornithoides mongoliensis
| `--Saurornithoides junior
`--Dromaeosauridae
|--Buitreraptor
|--Unenlagiinae
| |--Unenlagia
| `--Rahonavis
|--Microraptorinae
| |--Microraptor
| `--Sinornithosaurus
`--+--Velociraptorinae
| |--Velociraptor
| |--IGM 100/1015
| `--Deinonychus
`--Dromaeosaurinae
|--Saurornitholestes
|--Adasaurus
|--Achillobator
|--Utahraptor
`--Dromaeosaurus
In cases like this, where Tyrannosaurus, Shuvuuia, Therizinosaurus,
Oviraptor and Troodon are at
least as far from Ornithomimus as Passer is, Ornithomimosauria should have
priority over
Ornithomimiformes.
Clark et al., 2002
...continued from non-paravian tree above
`--Paraves
Eumaniraptora
|--+--Rahonavis
| `--Aves
| |--Archaeopterygidae
| | Archaeopteryx
| `--Ornithurae
| Confuciusornithidae
| Confuciusornis (+Passer?)
`--Deinonychosauria
|--Troodontidae
| |--Sinovenator
| `--+--Sinornithoides
| |--Byronosaurus
| `--+--Troodon
| `--+--Saurornithoides mongoliensis
| `--Saurornithoides junior
`--Dromaeosauridae
|--Sinornithosaurus
`--+--Microraptorinae
| Microraptor
`--+--Unenlagiinae
| Unenlagia
|--Velociraptorinae
| Velociraptor
|--IGM 100/1015
|--Deinonychus
|--Saurornitholestes
|--Adasaurus
|--Achillobator
|--Utahraptor
`--Dromaeosaurinae
Dromaeosaurus
Hwang et al., 2002
... continued from non-dromaeosaurid tree above
`--Dromaeosauridae
|--Microraptorinae
| Microraptor
`--+--Sinornithosaurus
`--+--Unenlagiinae
| Unenlagia
|--Velociraptorinae
| Velociraptor
|--IGM 100/1015
|--Deinonychus
|--Saurornitholestes
|--Adasaurus
|--Achillobator
|--Utahraptor
`--Dromaeosaurinae
Dromaeosaurus
Xu et al., 2002
... continued from non-maniraptoran main TWG tree
`--Maniraptora
|--Ornitholestes
`--+--Alvarezsauridae
| |--Alvarezsaurus
| `--+--Patagonykus
| `--Mononykinae
| |--Shuvuuia
| `--Mononykus
|--Oviraptoriformes
| |--Therizinosauria
| | |--Erlikosaurus (+Therizinosaurus?)
| | |--Segnosaurus
| | `--Alxasaurus
| `--Oviraptorosauria
| |--Incisivosaurus
| `--+--Caudipteryx
| `--Caenagnathoidea
| Oviraptorinae
| Ingeniinae
| |--Avimimus
| |--Caenagnathidae
| | Chirostenotes
| |--Microvenator
| |--Oviraptoridae
| | Oviraptor
| |--Ingenia
| |--Citipati
| `--+--Conchoraptor
| `--Rinchenia
`--Paraves
Eumaniraptora
|--Aves
| |--Rahonavis
| |--Archaeopterygidae
| | Archaeopteryx
| `--Ornithurae
| Confuciusornithidae
| Confuciusornis (+Passer?)
`--Deinonychosauria
|--Troodontidae
| |--Sinovenator
| `--+--Sinornithoides
| |--Byronosaurus
| `--+--Troodon
| `--+--Saurornithoides mongoliensis
| `--Saurornithoides junior
`--Dromaeosauridae
|--Sinornithosaurus
`--+--Unenlagiinae
| Unenlagia
|--Velociraptorinae
| Velociraptor
|--IGM 100/1015
|--Deinonychus
|--Saurornitholestes
|--Adasaurus
|--Achillobator
|--Utahraptor
`--Dromaeosaurinae
Dromaeosaurus
Ji et al., 2003
...continued from non-oviraptorosaurian main TWG tree
| `--Oviraptorosauria
| |--Caudipteryx
| `--Caenagnathoidea
| |--Caenagnathidae
| | |--Avimimus
| | |--Chirostenotes
| | `--Microvenator
| `--Oviraptoridae
| Oviraptorinae
| Ingeniinae
| |--Oviraptor
| |--Rinchenia
| |--Citipati
| |--Conchoraptor
| `--Ingenia
Norell et al., 2001
Neotetanurae
|--Sinraptoridae
| Sinraptor
|--Allosauroidea
| Allosauridae
| Allosaurus
`--Coelurosauria
Tyrannoraptora
|--Tyrannosauroidea
| Tyrannosauridae
| |--Gorgosaurus
| `--Tyrannosaurinae
| Tyrannosaurus
`--Maniraptoriformes
|--Ornithomimiformes
| Ornithomimosauria
| |--Pelecanimimus
| |--Harpymimus
| |--Garudimimus
| `--Ornithomimidae
| |--Gallimimus
| `--Struthiomimus (+Ornithomimus)
`--Maniraptora
|--Ornitholestes
`--+--Alvarezsauridae
| |--Alvarezsaurus
| `--+--Patagonykus
| `--Mononykinae
| |--Shuvuuia
| `--Mononykus
`--+--Oviraptoriformes
| Troodontidae
| |--+--Sinornithoides
| | |--Byronosaurus
| | `--+--Troodon
| | `--+--Saurornithoides mongoliensis
| | `--Saurornithoides junior
| `--+--Therizinosauria
| | |--Alxasaurus
| | |--Erlikosaurus (+Therizinosaurus?)
| | `--Segnosaurus
| `--Oviraptorosauria
| Caenagnathoidea
| |--Caenagnathidae
| | |--Avimimus
| | `--Chirostenotes
| `--Oviraptoridae
| |--Microvenator
| `--+--Caudipteryx
| `--Oviraptorinae
| Ingeniinae
| |--Oviraptor
| |--Rinchenia
| |--Citipati
| |--Conchoraptor
| `--Ingenia
`--Paraves
Eumaniraptora
|--+--Rahonavis
| `--Aves
| |--Archaeopterygidae
| | Archaeopteryx
| `--Ornithurae
| Confuciusornithidae
| Confuciusornis (+Passer?)
|--Unenlagiinae
| Unenlagia
|--Sinornithosaurus
|--Velociraptorinae
| Velociraptor
|--IGM 100/1015
|--Deinonychus
|--Saurornitholestes
|--Adasaurus
|--Achillobator
|--Utahraptor
`--Dromaeosauridae
Dromaeosaurinae
Dromaeosaurus
Another tree showing Troodontidae needs Oviraptor as an external specifier.
Hwang et al., 2004
Neotetanurae
|--Sinraptoridae
| Sinraptor
|--Allosauroidea
| Allosauridae
| Allosaurus
`--Coelurosauria
Tyrannoraptora
|--Tyrannosauroidea
| Tyrannosauridae
| |--Gorgosaurus
| `--Tyrannosaurinae
| Tyrannosaurus
`--Maniraptoriformes
|--Ornithomimiformes
| Ornithomimosauria
| |--Pelecanimimus
| `--+--Shenzhousaurus
| `--+--Harpymimus
| `--+--Garudimimus
| `--Ornithomimidae
| |--Archaeornithomimus
| |--Gallimimus
| |--Struthiomimus
| |--Anserimimus
| `--Ornithomimus
`--Maniraptora
|--Ornitholestes
`--+--Compsognathidae
| |--Huaxiagnathus
| `--+--Sinosauropteryx
| `--Compsognathus
`--+--Alvarezsauridae
| |--Alvarezsaurus
| `--+--Patagonykus
| `--Mononykinae
| |--Shuvuuia
| `--Mononykus
|--Oviraptoriformes
| |--Therizinosauria
| | |--Erlikosaurus (+Therizinosaurus?)
| | |--Segnosaurus
| | `--Alxasaurus
| `--Oviraptorosauria
| |--Incisivosaurus
| `--+--Caudipteryx
| `--Caenagnathoidea
| |--Caenagnathidae
| | |--Avimimus
| | |--Chirostenotes
| | `--Microvenator
| `--Oviraptoridae
| Oviraptorinae
| Ingeniinae
| |--Oviraptor
| |--Rinchenia
| |--Citipati
| |--Conchoraptor
| `--Ingenia
`--Paraves
Eumaniraptora
|--+--Rahonavis
| `--Aves
| |--Archaeopterygidae
| | Archaeopteryx
| `--Ornithurae
| Confuciusornithidae
| Confuciusornis (+Passer?)
`--Deinonychosauria
|--Troodontidae
| |--Sinovenator
| `--+--IGM 100/44
| `--+--Sinornithoides
| |--Byronosaurus
| `--+--Troodon
| `--+--Saurornithoides mongoliensis
| `--Saurornithoides junior
`--Dromaeosauridae
|--Microraptorinae
| Microraptor
|--Sinornithosaurus
`--+--Unenlagiinae
| Unenlagia
|--Velociraptorinae
| Velociraptor
|--IGM 100/1015
|--Deinonychus
|--Saurornitholestes
|--Adasaurus
|--Achillobator
|--Utahraptor
`--Dromaeosaurinae
Dromaeosaurus
Lu, 2004
... continued from non-maniraptoran tree above
`--Maniraptora
|--Ornitholestes
`--+--Compsognathidae
| |--Compsognathus
| |--Huaxiagnathus
| `--Sinosauropteryx
`--+--Oviraptoriformes
| |--Alvarezsauridae
| | |--Alvarezsaurus
| | `--+--Patagonykus
| | `--Mononykinae
| | |--Shuvuuia
| | `--Mononykus
| `--+--Therizinosauria
| | |--Erlikosaurus (+Therizinosaurus?)
| | |--Segnosaurus
| | `--Alxasaurus
| `--Oviraptorosauria
| |--Incisivosaurus
| `--+--Caudipteryx
| `--Caenagnathoidea
| Oviraptorinae
| Ingeniinae
| |--Nomingia
| |--Shixinggia
| |--Caenagnathidae
| | |--Chirostenotes
| | `--+--Avimimus
| | `--Microvenator
| |--NXMV
| |--Oviraptoridae
| | Oviraptor
| |--Rinchenia
| |--Citipati
| |--Nemegtia
| |--Khaan
| |--Conchoraptor
| |--Heyuannia
| `--Ingenia
`--Paraves
Eumaniraptora
Kirkland et al., 2005
Neotetanurae
|--Sinraptoridae
| Sinraptor
|--Allosauroidea
| Allosauridae
| Allosaurus
`--Coelurosauria
Tyrannoraptora
|--Tyrannosauroidea
| Tyrannosauridae
| |--Gorgosaurus
| `--Tyrannosaurinae
| Tyrannosaurus
`--Maniraptoriformes
|--Ornithomimiformes
| Ornithomimosauria
| |--Pelecanimimus
| `--+--Shenzhousaurus
| `--+--Harpymimus
| `--+--Garudimimus
| `--Ornithomimidae
| |--Archaeornithomimus
| `--+--+--Gallimimus
| | `--Struthiomimus
| `--+--Anserimimus
| `--Ornithomimus
`--Maniraptora
|--Ornitholestes
`--+--Compsognathidae
| |--Huaxiagnathus
| `--+--Compsognathus
| `--Sinosauropteryx
`--+--Oviraptoriformes
| |--Therizinosauria
| | |--Falcarius
| | `--Therizinosauroidea
| | Therizinosauridae
| | |--Beipiaosaurus
| | |--Alxasaurus
| | |--Nothronychus
| | |--Erlikosaurus (+Therizinosaurus?)
| | `--Segnosaurus
| `--Oviraptorosauria
| Caenagnatoidea
| |--Incisivosaurus
| |--Caudipteryx
| |--Avimimus
| |--Caenagnathidae
| | Chirostenotes
| |--Microvenator
| `--Oviraptoridae
| Oviraptorinae
| Ingeniinae
| |--Oviraptor
| |--Rinchenia
| |--Citipati
| |--Conchoraptor
| `--Ingenia
`--Paraves
Eumaniraptora
|--Alvarezsauridae
| |--Alvarezsaurus
| `--+--Patagonykus
| `--Mononykinae
| |--Shuvuuia
| `--Mononykus
|--+--Rahonavis
| `--Aves
| |--Archaeopterygidae
| | Archaeopteryx
| `--Ornithurae
| Confuciusornithidae
| Confuciusornis (+Passer?)
|--Troodontidae
| |--Sinovenator
| `--+--IGM 100/44
| `--+--Sinornithoides
| |--Byronosaurus
| `--+--Troodon
| `--+--Saurornithoides mongoliensis
| `--Saurornithoides junior
|--Microraptorinae
| Microraptor
|--Sinornithosaurus
`--Dromaeosauridae
|--Unenlagiinae
| Unenlagia
|--Velociraptorinae
| Velociraptor
|--IGM 100/1015
|--Deinonychus
|--Saurornitholestes
|--Adasaurus
|--Achillobator
|--Utahraptor
`--Dromaeosaurinae
Dromaeosaurus
Makovicky et al., 2005
...continued from non-dromaeosaurid main TWG tree
`--Dromaeosauridae
|--Unenlagiinae
| |--Buitreraptor
| `--+--Unenlagia + Neuquenraptor
| `--Rahonavis
`--+--Microraptorinae
| |--Microraptor
| `--Sinornithosaurus
`--+--Velociraptorinae
| |--Velociraptor
| |--IGM 100/1015
| `--Deinonychus
`--Dromaeosaurinae
|--Saurornitholestes
|--Adasaurus
|--Achillobator
|--Utahraptor
`--Dromaeosaurus
Mayr et al., 2005
... continued from non-paravian TWG tree
`--Paraves
Aves
|--Archaeopterygidae
| |--Rahonavis
| `--Archaeopteryx
`--Ornithurae
|--Troodontidae
| |--Sinovenator
| `--+--Byronosaurus
| `--+--Sinornithoides
| `--+--Troodon
| `--+--Saurornithoides mongoliensis
| `--Saurornithoides junior
`--Eumaniraptora
|--+--Microraptorinae
| | Microraptor
| `--Confuciusornithidae
| Confuciusornis (+Passer?)
`--Dromaeosauridae
|--Sinornithosaurus
`--+--Unenlagiinae
| Unenlagia
|--Velociraptorinae
| Velociraptor
|--IGM 100/1015
|--Deinonychus
|--Saurornitholestes
|--Adasaurus
|--Achillobator
|--Utahraptor
`--Dromaeosaurinae
Dromaeosaurus
Xu and Zhang, 2005
Neotetanurae
|--Allosauroidea
| Allosauridae
| Allosaurus
`--Coelurosauria
Tyrannoraptora
|--Tyrannosauroidea
| Tyrannosauridae
| |--Gorgosaurus
| `--Tyrannosaurinae
| Tyrannosaurus
`--Maniraptoriformes
|--Ornithomimiformes
| Ornithomimosauria
| |--Pelecanimimus
| |--Harpymimus
| |--Garudimimus
| |--Gallimimus
| `--Ornithomimidae
| Struthiomimus (+Ornithomimus?)
`--Maniraptora
|--Ornitholestes
`--+--Alvarezsauridae
| |--Alvarezsaurus
| `--+--Patagonykus
| `--Mononykinae
| |--Shuvuuia
| `--Mononykus
`--+--Oviraptoriformes
| |--Therizinosauria
| | |--Alxasaurus
| | `--+--Erlikosaurus (+Therizinosaurus?)
| | `--Segnosaurus
| `--Oviraptorosauria
| |--Incisivosaurus
| `--+--Caudipteryx
| `--Caenagnathoidea
| |--Caenagnathidae
| | |--Avimimus
| | `--Chirostenotes
| `--Oviraptoridae
| Ingeniinae
| |--Oviraptor
| |--Ingenia
| `--+--Microvenator
| `--+--Rinchenia
| `--Conchoraptor
`--Paraves
Aves
Eumaniraptora
|--Pedopenna
|--Epidendrosaurus
|--Archaeopterygidae
| Archaeopteryx
|--Rahonavis
|--Shenzhouraptor
|--Ornithurae
| Confuciusornithidae
| Confuciusornis (+Passer?)
`--Deinonychosauria
|--Troodontidae
| |--Sinovenator
| `--+--Byronosaurus
| `--+--Sinornithoides
| `--+--Troodon
| `--+--Saurornithoides mongoliensis
| `--Saurornithoides junior
`--Dromaeosauridae
|--Microraptorinae
| |--Microraptor
| |--Sinornithosaurus
| `--Graciliraptor
`--+--Unenlagiinae
| Unenlagia
|--Velociraptorinae
| Velociraptor
|--Deinonychus
|--Saurornitholestes
|--Adasaurus
|--Dromaeosaurinae
| Dromaeosaurus
|--Achillobator
`--Utahraptor
A possibility in this tree is (Passer (Deinonychus, Archaeopteryx)), which
puts Aves and
Eumaniraptora into conflict. I recommend Aves have priority in this case.
Allain, 2002
Neotetanurae
|--Allosauroidea
| |--Allosauridae
| | |--Allosaurus
| | `--Acrocanthosaurus
| `--+--Sinraptoridae
| | Sinraptor
| `--Carcharodontosauridae
| |--Giganotosaurus
| `--Carcharodontosaurus
`--Coelurosauria
Tyrannoraptora
|--Tyrannosaurus
`--+--Compsognathidae
| Compsognathus
`--Aves
Archaeopteryx (+Passer?)
Chiappe and Walker, 2002
After deletion of many taxa, to provide structure to tree.
|--Archaeopteryx
`--+--Eoenantiornis
|--Sinornis
|--Eoalulavis
|--Gobipteryx
`--+--Neuquenornis
|--Concornis
`--Enantiornis
Coria and Currie, 2002
Neotetanurae
Tyrannoraptora
|--Coelurosauria
| Eumaniraptora
| Dromaeosauridae (+Passer?)
|--Troodontidae
| |--Tyrannosauroidea
| | Tyrannosauridae
| `--Troodontidae
`--Allosauroidea
Abelisauridae
|--Abelisaurus
|--Carnotaurinae
| Carnotaurus
|--Allosauridae
| Allosaurus
|--Acrocanthosaurus
`--+--Sinraptoridae
| Sinraptor
`--Carcharodontosauridae
|--Giganotosaurus
`--Carcharodontosaurus
This indicates Tyrannosaurus should be an external specifier for
Troodontidae.
Gishlick, 2002
Neotetanurae
|--Allosauroidea
| Allosauridae
| Allosaurus
`--Coelurosauria
|--Alvarezsauridae
| Mononykus (+Shuvuuia?)
`--+--Coelurus
`--+--Compsognathidae
| Compsognathus
|--Scipionyx
|--Sinosauropteryx
`--+--Ornitholestes
`--+--Microvenator
`--+--Therizinosauria
| Beipiaosaurus (+Therizinosaurus)
|--Caudipteryx
`--+--Oviraptoriformes
| Oviraptorosauria
| Oviraptoridae
`--Paraves
|--Saurornitholestes
`--Eumaniraptora
Aves
|--Protarchaeopteryx
|--Dromaeosauridae
| Deinonychus (+Dromaeosaurus?)
|--Microraptorinae
| Sinornithosaurus (+Microraptor?)
|--Archaeopterygidae
| Archaeopteryx
`--Ornithurae
|--Confuciusornithidae
| Confuciusornis
`--Neornithes
Maryanska et al., 2002
After inclusion of Troodontidae, Mononykinae and Microvenator, which were
needlessly excluded
from the published analysis.
Neotetanurae
|--Allosauroidea
`--Coelurosauria
Tyrannoraptora
|--Tyrannosauroidea
| Tyrannosaurinae
`--Maniraptoriformes
|--Ornithomimiformes
| Ornithomimosauria
`--Maniraptora
Eumaniraptora
|--Deinonychosauria
| |--Dromaeosauridae
| | Velociraptorinae
| `--Troodontidae
`--+--Alvarezsauridae
| Mononykinae
`--+--Therizinosauria
| Therizinosauroidea
`--Aves
|--Archaeopterygidae
| Archaeopteryx
`--Ornithurae
|--Paraves
| Confuciusornithidae
| Confuciusornis (+Passer?)
`--Oviraptoriformes
Oviraptorosauria
|--Avimimus
`--+--Caudipteryx
`--Caenagnathoidea
|--Caenagnathidae
| |--Chirostenotes
| `--Nomingia
`--Oviraptoridae
Oviraptorinae
|--+--Microvenator
| `--Rinchenia (+Oviraptor?)
`--+--Citipati sp.
`--Ingeniinae
|--Conchoraptor
`--Ingenia
Lu, 2004
This was taken from the matrix, with characters ordered and unordered
properly, some
necessitating switching states 0 and 1 in all taxa.
Neotetanurae
Tyrannoraptora
|--Allosauroidea
| Tyrannosauroidea
| |--Allosauroidea sensu stricto
| `--Tyrannosaurinae
`--Coelurosauria
Maniraptoriformes
|--Ornithomimiformes
| Ornithomimosauria
`--Maniraptora
|--Paraves
| |--Alvarezsauridae
| | Mononykinae
| `--Eumaniraptora
| |--Troodontidae
| |--Dromaeosauridae
| | Velociraptorinae
| `--Aves
| Archaeopterygidae
| Archaeopteryx (+Passer?)
`--Oviraptoriformes
|--Therizinosauria
| Therizinosauroidea
`--Oviraptorosauria
|--Incisivosaurus
`--+--Caudipteryx
`--+--Avimimus
`--Caenagnathoidea
Oviraptorinae
|--Oviraptoridae
| Oviraptor
`--Caenagnathidae
|--+--Shixinggia
| `--+--Nomingia
| `--Chirostenotes
`--Ingeniinae
|--Khaan
|--Heyuannia
|--+--NXMV
| `--Ingenia
`--+--Conchoraptor
`--+--+--Citipati osmolskae
| `--Nemegtia
`--+--Citipati sp.
`--+--Rinchenia
`--Microvenator
Here's an interesting situation with Oviraptor's placement. The main
problem is Oviraptorinae,
which should be a stem from Ingenia, instead of a node with Citipati. This
would assure it a
logical placement in Maryanska et al.'s topology above as well.
Varricchio, 2002
Eumaniraptora
|--Dromaeosauridae
|--Troodontidae
`--Ornithothoraces
Carinatae
|--Enantiornithes (+Sinornis)
|--Ichthyornis
`--Euornithes
Neornithes
Currie et al., 2003 and Xu et al., 2004
Neotetanurae
|--Allosauroidea
| Allosauridae
| Allosaurus
`--Coelurosauria
|--Compsognathidae
| Sinosauropteryx (+Compsognathus)
`--Maniraptoriformes
|--Ornithomimiformes
| Ornithomimosauria
| Gallimimus (+Ornithomimus)
`--Maniraptora
Tyrannoraptora
|--Paraves
| Dromaeosauridae (+Passer?)
| |--Velociraptorinae
| | Velociraptor
| `--Dromaeosaurinae
| Dromaeosaurus
`--Oviraptoriformes
Troodontidae
|--Sinovenator (+Troodon)
`--Tyrannosauroidea
|--Oviraptorosauria
| Citipati (+Oviraptor)
`--+--Dilong
`--Tyrannosauridae
|--Albertosaurinae
| |--Albertosaurus
| `--Gorgosaurus
`--Tyrannosaurinae
|--Daspletosaurus
`--+--+--Alioramus
| `--Tarbosaurus
`--+--Nanotyrannus
`--Tyrannosaurus
Another case of Troodontidae needing Tyrannosaurus as an external specifier.
Tyrannosauroidea
also needs Oviraptor as an external specifier in this tree.
Kobayashi and Lu, 2003
|--Allosaurus
|--Tyrannosauroidea
| Tyrannosaurinae
`--Ornithomimosauria
|--Pelecanimimus
`--+--Harpymimus
`--+--Garudimimus
`--Ornithomimidae
|--Archaeornithomimus
`--+--+--Anserimimus
| `--Gallimimus
`--+--Struthiomimus
`--+--Ornithomimus
`--Dromiceiomimus
Clark et al., 2004
After deletion of Enigmosaurus, Nothronychus and Eshanosaurus to provide
structure to tree.
Maniraptoriformes
|--Ornithomimiformes
| Ornithomimosauria
|--Alvarezsauridae
|--Troodontidae
|--Maniraptora
| Paraves
| Eumaniraptora
| Dromaeosauridae (+Passer?)
`--Oviraptoriformes
|--Oviraptorosauria
`--Therizinosauria
Therizinosauroidea
|--Beipiaosaurus
`--+--Alxasaurus
`--Therizinosauridae
|--Neimongosaurus
|--Nanshiungosaurus
|--Erlikosaurus
|--Therizinosaurus
`--Segnosaurus
Currie and Varricchio, 2004
|--Allosaurus
`--Deinonychosauria
|--Troodontidae
`--Dromaeosauridae
|--Dromaeosaurinae
| Dromaeosaurus
`--Velociraptorinae
|--Saurornitholestes
`--+--Velociraptor
`--+--Bambiraptor
`--+--Atrociraptor
`--Deinonychus
Franzosa, 2004
Neotetanurae
|--Ceratosaurus
|--Majungatholus
|--Allosauroidea
| Allosauridae
| Allosaurus
|--Acrocanthosaurus
|--Carcharodontosauridae
| Carcharodontosaurus
`--Coelurosauria
Tyrannoraptora
|--Tyrannosauroidea
| Tyrannosaurus
`--Aves
|--Oviraptoriformes
| Oviraptorosauria
| Citipati (+Oviraptor)
|--Troodontidae
| Saurornithoides mongoliensis (+Troodon?)
|--Zos Canyon troodontid
|--Byronosaurus
|--Archaeopterygidae
| Archaeopteryx
`--Paraves
Ornithurae
Neornithes
Osmolska et al., 2004
Eumaniraptora
|--Dromaeosauridae
| Velociraptor (+Deinonychus)
`--+--Paraves
| Archaeopteryx (+Passer?)
`--Oviraptoriformes
Oviraptorosauria
|--Avimimus
`--+--Caudipteryx
`--Caenagnathoidea
|--Caenagnathidae
| Chirostenotes
`--Oviraptoridae
|--Oviraptorinae
| |--Rinchenia (+Oviraptor?)
| `--+--Citipati osmolskae
| `--Citipati sp.
`--Ingeniinae
|--Ingenia
`--+--Khaan
`--Conchoraptor
Senter et al., 2004
Neotetanurae
|--Allosauroidea
| Allosauridae
| Allosaurus
`--Coelurosauria
Tyrannoraptora
|--Tyrannosauroidea
| Tyrannosaurus
`--+--Ornitholestes
`--+--Compsognathidae
| Sinosauropteryx (+Compsognathus?)
`--Maniraptoriformes
|--Ornithomimiformes
| |--Ornithomimosauria
| | |--Pelecanimimus
| | `--Ornithomimidae
| | Struthiomimus (+Ornithomimus)
| `--Troodontidae
| |--Byronosaurus
| `--+--Sinornithoides
| `--+--Troodon
| |--Saurornithoides mongoliensis
| `--Saurornithoides junior
`--Maniraptora
|--Oviraptoriformes
| Oviraptorosauria
| |--Protarchaeopteryx + Incisivosaurus
| `--+--Caudipteryx
| `--Khaan (+Oviraptor)
`--Paraves
Eumaniraptora
|--Aves
| |--Archaeopterygidae
| | Archaeopteryx
| `--Ornithurae
| |--Rahonavis
| |--Shenzhouraptor
| `--Ornithothoraces
| |--Enantiornithes
| | Sinornis
| `--Euornithes
| Yanornis (+Passer?)
`--Dromaeosauridae
|--Sinovenator
`--+--Microraptorinae
| |--Bambiraptor
| `--+--Sinornithosaurus
| `--+--NGMC 91
| `--+--Microraptor zhaoianus
| |--Microraptor gui
| `--Cryptovolans
`--+--Velociraptorinae
| Velociraptor
`--Dromaeosaurinae
|--Adasaurus
`--+--Saurornitholestes
`--+--Deinonychus
`--+--Achillobator
`--+--Utahraptor
`--Dromaeosaurus
Carr et al., 2005
|--Eoraptor
`--+--Therizinosauria
| Herrerasauridae
| |--Herrerasaurus
| `--Erlikosaurus (+Therizinosaurus?)
`--Neotheropoda
|--Coelophysoidea
| Coelophysis
`--Neotetanurae
|--Allosauroidea
| Allosauridae
| Allosaurus
`--+--Yangchuanosaurus
`--+--Sinraptoridae
| Sinraptor
`--Tyrannoraptora
Maniraptoriformes
|--Carcharodontosauridae
| Acrocanthosaurus (+Carcharodontosaurus?)
|--Ornithomimiformes
| |--Ornithomimosauria
| | Ornithomimidae
| `--Troodontidae
| Saurornithoides (+Troodon)
|--Maniraptora
| Eumaniraptora
| Dromaeosauridae (+Passer?)
| |--Deinonychus (+Dromaeosaurus?)
| `--Velociraptor
`--Tyrannosauroidea
|--Dryptosaurus
`--+--Appalachiosaurus
`--Tyrannosauridae
|--Albertosaurinae
| |--Gorgosaurus
| `--Albertosaurus
`--Tyrannosaurinae
|--Daspletosaurus
`--+--Tarbosaurus
|--Shanshanosaurus
`--Tyrannosaurus
I think the placement of Erlikosaurus here is too unlikely to bother
justifying a definitional
adjustment for. The tree does justify Sereno's use of Passer as an external
specifier to
Sinraptoridae though.
Novas et al., 2005
Neotetanurae
|--Allosauroidea
| |--Monolophosaurus
| `--+--Sinraptoridae
| | Sinraptor
| `--+--Allosauridae
| | |--Allosaurus
| | `--Acrocanthosaurus
| `--Carcharodontosauridae
| |--Tyrannotitan
| `--+--Giganotosaurus
| `--Carcharodontosaurus
`--Coelurosauria
Tyrannoraptora
|--Tyrannosauroidea
| Tyrannosaurus
`--+--Oviraptoriformes
| Oviraptorosauria
| Oviraptoridae
`--Paraves
Eumaniraptora
Dromaeosauridae (+Passer?)
Rauhut and Xu, 2005
Neotetanurae
|--Allosauroidea
| Allosauridae
| Allosaurus
`--Coelurosauria
|--Tugulusaurus
`--+--Coelurus
|--Compsognathidae
`--Tyrannoraptora
Maniraptoriformes
|--Tyrannosauroidea
|--Nqwebasaurus
|--Ornitholestes
|--Ornithomimiformes
| Ornithomimosauria
`--Maniraptora
|--Oviraptoriformes
| |--Therizinosauria
| | Therizinosauroidea
| `--Oviraptorosauria
`--Paraves
|--+--Bagaraatan
| `--Xinjiangovenator
`--Eumaniraptora
|--Aves
`--Deinonychosauria
|--Dromaeosauridae
`--Troodontidae
Holtz et al., 2004 and Holtz, 2004
These derive from the same matrix.
Neotetanurae
|--Allosauroidea
| |--Lourinhanosaurus
| `--+--Monolophosaurus
| `--+--Sinraptoridae
| `--+--Allosauridae
| | Allosaurus
| `--Carcharodontosauridae
| |--Neovenator
| `--+--Acrocanthosaurus
| `--+--Carcharodontosaurus
| `--Giganotosaurus
`--Coelurosauria
|--+--Fukuiraptor
| `--Siamotyrannus
`--+--Compsognathidae
| |--Nqwebasaurus
| `--+--Sinosauropteryx
| `--Compsognathus
`--+--Scipionyx
`--+--+--Xuanhanosaurus
| `--+--Ornitholestes
| `--Proceratosaurus
`--Tyrannoraptora
|--Tyrannosauroidea
| |--Bagaraatan
| `--+--Stokesosaurus
| |--Dryptosaurus
| `--+--Eotyrannus
| `--Tyrannosauridae
| |--Appalachiosaurus
| |--Albertosaurinae
| | Albertosaurus
| |--Gorgosaurus
| `--Tyrannosaurinae
| |--Daspletosaurus
| `--+--Tarbosaurus
| `--Tyrannosaurus
`--+--Coelurus
`--Maniraptoriformes
|--Ornithomimiformes
| Ornithomimosauria
| |--Harpymimus
| `--+--Pelecanimimus
| `--+--Garudimimus
| `--Ornithomimidae
`--Maniraptora
|--Oviraptoriformes
| |--Therizinosauria
| | Therizinosauroidea
| | Therizinosauridae
| | |--Nothronychus
| | `--+--Erlikosaurus (+Therizinosaurus?)
| | `--Beipiaosaurus
| `--Oviraptorosauria
| |--Caudipteryx
| `--Caenagnathoidea
| |--Caenagnathidae
| | Chirostenotes
| `--Oviraptoridae
| |--Khaan
| `--Oviraptor
`--Paraves
|--Alvarezsauridae
| |--Alvarezsaurus
| `--+--Patagonykus
| `--Mononykinae
`--Eumaniraptora
|--Protarchaeopteryx
|--Aves
| |--Archaeopterygidae
| | Archaeopteryx
| `--Ornithurae
| |--Rahonavis
| `--+--Confuciusornithidae
| | Confuciusornis
| `--Ornithothoraces
`--Deinonychosauria
|--Troodontidae
| |--Sinovenator
| `--+--Sinornithoides
| `--Saurornithoides + Troodon
`--Dromaeosauridae
|--Bambiraptor
`--+--Sinornithosaurus
`--+--Velociraptorinae
| Velociraptor
`--Dromaeosaurinae
|--Deinonychus
`--Dromaeosaurus
Here's Ornithomimiformes vs. Ornithomimosauria again. Also, in this tree
Therizinosauridae
equals Therizinosauroidea, due to the (Nothronychus (Therizinosaurus,
Beipiaosaurus)) topology,
assuming Therizinosaurus is closest to Erlikosaurus. While unlikely, I
suggest Therizinosauridae
have priority based on the publishing dates.
Mortimer, in prep.
And here's my latest tree, a 90% consensus, leaving out some taxa to limit
the amount of
polytomies. Note I don't support everything in this tree, but it's
interesting to see how
Sereno's definitions would apply.
|--Monolophosaurus
`--+--+--Fukuiraptor
| |--+--Labocania
| | `--"Alashansaurus"
| `--+--Elmisaurus
| `--+--Megaraptor
| `--Tanycolagreus
`--Neotetanurae
|--Allosauroidea
| |--+--Sinraptoridae
| | | Sinraptor
| | `--Allosauridae
| | Allosaurus
| `--Carcharodontosauridae
| |--Acrocanthosaurus (+Carcharodontosaurus?)
| `--Dryptosaurus
`--Coelurosauria
Tyrannoraptora
|--Tyrannosauroidea
| |--+--Dilong
| | `--Eotyrannus
| `--+--Appalachiosaurus
| `--Tyrannosauridae
| |--Albertosaurinae
| | Albertosaurus
| `--+--Gorgosaurus
| `--Tyrannosaurinae
| |--Daspletosaurus
| `--+--Tarbosaurus
| `--+--Bagaraatan
| `--+--Aviatyrannis
| `--Tyrannosaurus
`--+--+--Huaxiagnathus
| `--Sinosauropteryx
`--+--Coelurus
`--+--+--Ornitholestes
| `--Nqwebasaurus
|--NGMC 2124
`--+--Proceratosaurus
`--+--Compsognathidae
| Compsognathus
`--+--Borsti
`--+--Scipionyx
`--Maniraptoriformes
|--Ornithomimiformes
| Ornithomimosauria
| |--+--Shenzhousaurus
| | `--"Grusimimus"
| `--+--+--Garudimimus
| | `--+--Harpymimus
| | `--Anserimimus
| `--Ornithomimidae
| |--Sinornithomimus
| `--+--Archaeornithomimus
| `--derived ornithomimids
`--Maniraptora
|--Alvarezsauridae
| |--Archaeornithoides
| `--+--Pelecanimimus
| `--+--Patagonykus
| `--+--Alvarezsaurus
| `--Mononykinae
| |--Mononykus
| `--+--Parvicursor
| `--Shuvuuia
`--Therizinosauroidea
Therizinosauridae
|--Therizinosauria
| Therizinosaurus
`--+--+--Nothronychus
| `--+--Alxasaurus
| `--+--+--Erlianosaurus
| | `--Neimongosaurus
| `--+--IVPP V11309
| `--+--+--Beipiaosaurus
| | `--Erlikosaurus
| `--+--Segnosaurus
|
`--Nanshiungosaurus
Caenagnathoidea
|--Oviraptoriformes
| Oviraptorosauria
| Oviraptoridae
| |--+--Caudipteryx
| | `--Nomingia
| `--+--+--Caenagnathus collinsi
| | `--CMNH caenagnathid
| `--+--+--Nemegtia
| | `--Heyuannia
| `--+--Shixinggia
| `--Oviraptorinae
| |--+--Microvenator
| | `--Oviraptor
| `--+--+--RTMP 79.20.1
| | `--+--Deinocheirus
| | `--Citipati
| `--Ingeniinae
| |--+--Rinchenia
| | `--+--Ingenia
| | `--NXMV
| `--+--Khaan
| `--+--Conchoraptor
| `--ZPAL MgD/I-95
`--Paraves
Caenagnathidae
|--+--ROM 43250
| `--+--Avimimus
| `--+--Incisivosaurus
| `--Falcarius
`--+--Protarchaeopteryx
`--+--WDC DML001
`--+--+--Yixianosaurus
| `--Unenlagia paynemili
`--+--Chirostenotes holotype
`--+--Ukhaa Tolgod troodontid
`--Eumaniraptora
|--Dromaeosauridae
| |--Richardoestesia
| `--+--Sinornithosaurus
| `--+--Microraptorinae
| | |--Bambiraptor
| | `--+--+--Graciliraptor
| | | `--NGMC 91
| | `--+--Microraptor gui
| | `--+--Microraptor zhaoianus
| | `--Cryptovolans
| `--+--Saurornitholestes
| `--+--Velociraptorinae
| | |--IGM 100/1015
| | `--+--Deinonychus
| | `--+--Velociraptor
| | `--Achillobator
| `--Dromaeosaurinae
| |--+--Utahraptor
| | `--Atrociraptor
| `--+--Dromaeosaurus
| `--Adasaurus
`--+--Unenlagiinae
| Unenlagia
`--+--Troodontidae
| |--Sinovenator
| `--+--Sinusonasus
| `--+--Byronosaurus
| `--+--IGM 100/44
| `--+--Sinornithoides
| `--+--Troodon
| `--+--Saurornithoides
mongoliensis
| `--Saurornithoides junior
`--+--Mei
`--Aves
|--Archaeopterygidae
| |--+--Buitreraptor
| | `--Rahonavis
| `--+--+--Jinfengopteryx
| | `--Scansoriopterygidae
| `--+--Archaeopteryx
| `--Wellnhoferia
`--Ornithurae
|--+--Sapeornis
| `--Omnivoropteryx
`--+--Yandangornis
`--+--Shenzhouraptor
`--+--Confuciusornithidae
| |--Confuciusornis
| `--Changchengornis
`--+--Jixiangornis
`--+--Gobipteryx
Ornithothoraces
|--Enantiornithes
| |--Concornis
| |--Enantiornis
| `--+--+--Eoenantiornis
| | `--LP-4450-IEI
| `--+--+--Eocathayornis
| | `--+--Aberratiodontus
| | `--Jibeinia
| `--+--Sinornis
| `--+--Vescornis
| `--+--Longirostravis
| `--+--Longipteryx
| `--Protopteryx
`--Euornithes
|--+--Eoalulavis
| `--Liaoningornis
`--+--Neuquenornis
`--+--Iberomesornis
`--+--Patagopteryx
`--+--Songlingornis
`--+--Yanornis
`--+--+--Yixianornis
| `--+--Apsaravis
| `--Ambiortus
`--+--Boluochia
`--Carinatae
|--Ichthyornis
`--+--Hesperornithes
`--+--Iaceornis (+Passer?)
`--+--Hongshanornis
`--"Archaeorhynchus"
There are some issues with Therizinosauroidea and Therizinosauridae since my
matrix refuses to
place Therizinosaurus with segnosaurs. Similarly, since Chirostenotes is a
basal paravian in
this tree, Caenagnathoidea, Caenagnathidae and Oviraptoridae are
compromised. But if
Caenagnathus were used as the specifier for these clades, normal composition
would be maintained.
Note a stem-based Oviraptorinae would also keep its composition more
logical. In this tree,
Sereno's use of Sinornis as an internal specifier for Ornithothoraces
(instead of Iberomesornis)
leads to better compositional stability.
Summary of conclusions to follow...
Mickey Mortimer