[Date Prev][Date Next][Thread Prev][Thread Next][Date Index][Thread Index][Subject Index][Author Index]

The Dinosauria 2nd Ed. - Saurischia taxonomy and phylogeny

First, some synonymizations which haven't been otherwise officially listed
in the literature yet.
Caseosaurus crosbyensis = Chindesaurus bryansmalli
Edmarka rex = Torvosaurus tanneri
Dromiceiomimus brevitertius, D. samueli = Ornithomimus edmontonicus
- No statistical support for supposed differences in ratios between taxa,
shares bifurcated dorsal quadratojugal process with Ornithomimus.  Suppose
this means the TWG's Ornithomimus OTU includes Dromiceiomimus data.
Holbotia ponomarenkoi = Ambiortus dementjevi
- Not true (Kurochkin, 2001).
Cetiosaurus leedsi = Cetiosauriscus stewarti
- The former is a titanosauriform and was named prior to the latter (a
diplodocoid?), so this is not true.
Eobrontosaurus yahnahpin = Camarasaurus sp.
- Long cervical ribs and distally flared scapula are unlike diplodocoids.
Cervical vertebrae, scapulacoracoid and humerus almost indistinguishable
from Camarasaurus.

Next, the phylogenies presented in each chapter.  These are each based on a
cladistic analysis whose data matrix is still unavailable, though they
should be on http://dinosauria.ucpress.edu/ sometime.  I've presented the
phylogenies as an analysis-based backbone with non-analyzed taxa inserted
where the authors classify them.  It follows this format-
 |(G. species1)
 `(G. species2)
Alpha, Beta and genus are taxa which were in the data matrix.  Beta is
Cladia incertae sedis.  Gamma, Delta and Epsilon weren't analyzed.  Delta is
only questionably in its position, while Epsilon is Cladia incertae sedis.
Finally, the above indicates the genus was an OTU, but each species wasn't
included separately.  Nomina dubia are listed under 'nd'.

9 taxa, 107 chars, CI .5914
   |(*unnamed herrerasaurian of Galton, 1985)
   |  `--other Ornithischia
      |  |(?Chindesaurus)
      |  `--Herrerasauridae
      |     |--Herrerasaurus
      |     `--Staurikosaurus
            |  `--Sauropodomorpha
               `--other Theropoda
nd- Agnostiphys, "Teyuwasu", Thecospondylus
This is from his SVP 2001 talk, and he comments on the affinities of
excluded taxa in more depth than most authors.  Alwalkeria lacks some
theropod characters; Chindesaurus has a sauropodomorph character, but can be
excluded from Theropoda, Sauropodomorpha or Herrerasauridae; Aliwalia
resembles sauropodomorphs more than theropods, but cannot even be safely
assigned to Saurischia; Agnostiphys is dinosaurian, possibly eusaurischian,
but not theropodan, and resembles Herrerasaurus and sauropodomorphs in
astragalar anatomy; Saltopus has saurischian characters, but lacks
dinosaurian characters; Spondylosoma resembles herrerasaurids, but also has
non-dinosaurian characters.  It takes 14 more steps to place herrerasaurids
outside Dinosauria (Eoraptor and Guaibasaurus stay as saurischians), and 2
more steps to follow Sereno's topology (Guaibasaurus stays as a theropod).

Tykoski and Rowe
19 taxa, 175 chars, CI .588
         |  |--Allosaurus
         |  `--+--Torvosaurus
         |     `--Spinosauridae
            |  |(*Camposaurus)
            |  |(*"Liliensternus" airelensis)
            |  |(*Podokesaurus)
            |  |(*shake-n-bake theropod)
            |  |--Dilophosaurus
            |  `--+--Liliensternus
            |     |(*Gojirasaurus)
            |     `--+--Procompsognathus
            |        `--Coelophysidae
            |           |*-Segisaurus
            |           |--Coelophysis
            |           `--+--Syntarsus rhodesiensis
            |              `--Syntarsus kayentakatae
                  |  |(C. nasicornis)
                  |  |(C. ?dentisulcatus)
                  |  `(C. ?magnicornis)
                     |  |--Noasaurus
                     |  |(Velocisaurus)
                     |  |(Laevisuchus)
                     |  `--Masiakasaurus
                        |  Abelisaurus
nd- Ceratosaurus roechlingi, Coeluroides, Coelurus longicollis,
Dolichosuchus, Dryptosauroides, Eucoelophysis, Genyodectes, Halticosaurus,
Jubbulpuria, Labrosaurus stechowi, Labrosaurus sulcatus, Lametasaurus,
Longosaurus, Ornithomimoides, Ornithomimoides barasimilis, Orthogoniosaurus,
Pterospondylus, Sarcosaurus andrewsi, Tanystrosuchus, Tanystropheus
willistoni, Velocipes
The authors suggest Ceratosaurus nasicornis' holotype is adult, perhaps
indicating Madsen and Welles' species are valid.  They claim that
Liliensternus has no apomorphies, and subsequently don't refer L. airelensis
to the genus, but Rauhut (2003) identified one.  It's quite possible they
think Aucasaurus, Indosuchus, Indosaurus and Xenotarsosaurus are
abelisaurids, though they never explicitly make this clear.

Holtz et al.
75 taxa, 638 chars, CI .354
      |(*Chilantaisaurus tashuikouensis)
      |(*Chilantaisaurus? sibiricus)
      |(*Erectopus superbus)
      |(*Erectopus sauvagei)
      |(*"Megalosaurus" hesperis)
      |(*"Zanclodon" cambrensis)
      |--"Szechuanosaurus" zigongensis
            |  |--Spinosauridae
            |  |  |--Baryonychinae
            |  |  |  |(Baryonyx)
            |  |  |  |(Suchomimus)
            |  |  |  `(Cristatusaurus)
            |  |  `--Spinosaurinae
            |  |     |--Spinosaurus
            |  |     |  |(S. aegyptiacus)
            |  |     |  `(S. maroccanus)
            |  |     |(Angaturuama)
            |  |     `--Irritator
            |  `--Megalosauridae
            |     |--Megalosaurinae
            |     |  |--Torvosaurus
            |     |  `--+--Megalosaurus
            |     |     `--Poekilopleuron
            |     `--Eustreptospondylinae
            |        |--Piatnitzkysaurus
            |        `--+--Eustreptospondylus
            |           `--+--Afrovenator
            |              `--"Poekilopleuron" valesdunensis
               |  |--Monolophosaurus
               |  |(Cryolophosaurus)
               |  |--+--Fukuiraptor
               |  |  `--Siamotyrannus
               |  |--Sinraptoridae
               |  |  |(Sinraptor dongi)
               |  |  |(Sinraptor hepingensis)
               |  |  |(Yangchuanosaurus hepingensis)
               |  |  `(Yangchuanosaurus magnus)
               |  |--Lourinhanosaurus
               |  |--Allosauridae
               |  |  |--Allosaurus
               |  |  |  |(A. fragilis)
               |  |  |  |(unnamed A. sp. of Chure, in press)
               |  |  |  `(A? tendagurensis)
               |  |  |(Saurophaganax)
               |  |  `(unnamed allosauroid of Perez-Moreno et al., 1993)
               |  `--Carcharodontosauridae
               |     |(*unnamed carcharodontosaurid of Alcober et al., 1998)
               |     |(*unnamed carcharodontosaurid of Coria and Currie,
               |     |--Neovenator
               |     `--+--Acrocanthosaurus
               |        `--+--Giganotosaurus
               |           `--Carcharodontosaurus
                  |(*Richardoestesia [as Ricardoestesia] gilmorei)
                  |(*Richardoestesia isosceles)
                  |(*unnamed Richardoestesia sp. from Milk River Formation)
                  |  |(Aristosuchus)
                  |  |(unnamed compsognathid of Martill et al., 2000)
                  |  |--Compsognathus
                  |  `--Sinosauropteryx
                        |  |--Ornitholestes
                        |  `--Nqwebasaurus
                                    |  `--Oviraptorosauria
                                          |  `--Dromaeosauridae
nd- Allosaurus medius, Altispinax, Aublysodon lateralis, Betasuchus,
Calamosaurus[as Calamospondylus foxi], Calamospondylus, Chienkosaurus,
Chuandongocoelurus, Coelurus gracilis, Creosaurus potens, Dandakosaurus,
Embasaurus, Inosaurus, Laelaps macropus, Megalosaurus chubutensis,
Megalosaurus hungaricus, Megalosaurus inexpectatus, Megalosaurus ingens,
Megalosaurus insignis, Megalosaurus lonzeensis, Megalosurus lydekkeri,
Megalosaurus meriani, Megalosaurus pannoniensis, Megalosaurus pombali,
Megalosaurus terquemi, Nuthetes, Phaedrolosaurus, Poekilopleuron schmidti,
Prodeinodon kwangshiensis, Prodeinodon, Scrotum, Siamosaurus, Sinocoelurus,
Streptospondylus cuvieri, Szechuanosaurus, Szechuanosaurus "yandonensis",
Teinurosaurus, Wakinosaurus, Walgettosuchus
Holtz et al. do an excellent job summarizing prior opinions of relationships
for many of the incertae sedis taxa here.

75 taxa, 638 chars, CI .354
   |  `--Maniraptora
      |(*"Ornithomimus" grandis)
      |(?*"Chilantaisaurus" maortuensis)
                  |  |--unnamed tyrannosaurid of Carr et al. in press
                  |  `--+--Albertosaurus
                  |     `--Gorgosaurus
                     |  |(D. torosus)
                     |  `(unnamed sp. from Lower Kirtland Formation)
nd- Aublysodon amplus, Aublysodon cristatus, Aublysodon lateralis,
Aublysodon, Deinodon, Laelaps macropus, Diplotomodon, Tyrannosaurus
"lanpingensis", Tyrannosaurus turpanensis
This is part of the tree shown in the Basal Tetanurae chapter, generated by
Holtz's dataset.  Alectrosaurus falls more derived than Eotyrannus, but
outside Alberto+Gorgo and Daspleto+Tarbo+Tyranno.

Makovicky et al.
50 taxa, 220 chars, CI .45
            |  |(O. velox)
            |  `--O. edmontonicus
nd- Archaeornithomimus bissektensis, Coelosaurus affinis, Coelosaurus,
Elaphrosaurus gautieri?, Elaphrosaurus iguidiensis?, Ornithomimus sedens,
Ornithomimus tenuis, Timimus
The authors exclude Deinocheirus because it lacks a supragenloid butress
flange, subglenoid shelf, closely appressed radius and ulna, and distally
placed flexor tubercles, though they say it may be a basal member of the
clade.  This, and the Troodontidae and Dromaeosauridae analyses are just the
TWG analysis (prior to the addition of Huaxiagnathus and Mei).  I don't know
why they didn't use Kobayashi's, which actually resolves ornithomimid

Clark et al.
10 taxa, 40 chars, CI .746
      |(Nanshiungosaurus? bohlini)
      |--Nanshiungosaurus brevispinus
nd- Chilantaisaurus zheziangensis
They actually included Nothronychus in the matrix, but excluded it from the
final run and never mention where it ends up.

Osmolska et al.
9 taxa, 161 chars, CI .689
         |  |(C. zoui)
         |  `(C. dongi)
            |  |--Chirostenotes
            |  |  |(C. pergracilis)
            |  |  `(C. elegans)
            |  |(?Nomingia)
            |  `(Caenagnathasia)
               |  |(*Oviraptor)
               |  |(*Heyuannia)
               |  |--Rinchenia
               |  `--+--IGM 100/42
               |     `--Citipati
nd- Thecocoelurus
Oviraptor, Microvenator and Nomingia were excluded from the analysis because
of the large amount of missing data and/or poor preservation.  They agree
with Maryanska et al. (2002) in questioning Elmisaurus' assignment to
Caenagnathidae, but never say what else it's supposed to be (it's listed as
'possible Oviraptorosauria').  Oddly, elegans doesn't follow rarus.  Reasons
for placing Heyuannia in Oviraptorinae are not given, and Oviraptor is said
to be hard to diagnose.

Makovicky and Norell
50 taxa, 220 chars, CI .45
   `--+--IGM 100/44
            `--+--Saurornithoides mongoliensis
               `--Saurornithoides junior
nd- Bradycneme, Elopteryx, Euronychodon, Euronychodon asiaticus,
Heptasteornis, Koparion, Laelaps cristatus, Laelaps explanatus, Laelaps
laevifrons, Paronychodon, Pectinodon, Tripriodon, Troodon asiamericanus,
Troodon isfarensis, Zapsalis

Norell and Makovicky
50 taxa, 220 chars, CI .45
|  |--M. zhaoianus
|  `(M. gui)
Problematic taxa- Euronychodon, Euronychodon asiaticus, Hulsanpes,
Megaraptor, Ornithodesmus, Pyroraptor, Unquillosaurus, Variraptor
nd- "Palaeopteryx"
Hulsanpes is said to lack dromaeosaurid characters, while Ornithodesmus'
affinities are "ambiguous" despite them listing nothing but support for a
dromaeosaurid identity.  Megaraptor is never discussed.

(uses Chiappe, 2001)
|  |(*Parvicursor)
|  |--Alvarezsaurus
|  `--+--Patagonykus
|     `--+--Shuvuuia
|        `--Mononykus
   |  |--Archaeopteryx
   |  |  |(A. lithographica)
   |  |  `(A. bavarica)
   |  `(Wellnhoferia)
         |  |--Changchengornis
         |  `--Confuciusornis
         |     |(C. sanctus)
         |     `(C. dui)
               |  |*(Alexornis)
               |  |*(Avisaurus archibaldi)
               |  |*(Avisaurus gloriae)
               |  |*(Boluochia)
               |  |*(Cathayornis)
               |  |*(Eocathayornis)
               |  |*(Eoenantiornis)
               |  |*(Enantiornis leali)
               |  |*(Enantiornis martini)
               |  |*(Enantiornis walkeri)
               |  |*(Gurilynia)
               |  |*(Halimornis)
               |  |*(Kizylkumavis)
               |  |*(Lectavis)
               |  |*(Lenesornis)
               |  |*(Nanantius)
               |  |*(Otogornis)
               |  |(?Protopteryx)
               |  |*(Sazavis)
               |  |*(Soroavisaurus)
               |  |*(Yungavolucris)
               |  |*(Zhyraornis kaskarovi)
               |  |*(Zhyraornis logunovi)
               |  |--+--Iberomesornis
               |  |  `--Noguerornis
               |  `--Euenantiornithes
               |     |--Concornis
               |     |--Eoalulavis
               |     |--Gobipteryx
               |     |--Neuquenornis
               |     `--Sinornis
                     |  |(Asiahesperornis)
                     |  |(Baptornis)
                     |  |(Coniornis)
                     |  |(Enaliornis barretti)
                     |  |(Enaliornis sedgwicki)
                     |  |--Hesperornis
                     |  |  |(H. regalis)
                     |  |  |(H. crassipes)
                     |  |  `(H. gracilis)
                     |  |(Judinornis)
                     |  |(Parahesperornis)
                     |  |(Pasquiaornis hardiei)
                     |  |(Pasquiaornis tankei)
                     |  `(Potamornis)
                           |  |--Ichthyornis
                           |  |("Graculavus" lentus)
                           |  `("Ichthyornis" tener)
nd- Ichthyornis minusculus, Ilerdopteryx, Ornithomimus minutus, Platanavis,
Protoavis?, Wyleyia?
For all the negative things I've said about this chapter, Padian gets credit
for placing Jibeinia between confuciusornithids and ornithothoraceans.  This
chapter doesn't get its own matrix, Padian merely refers the viewer to
Chiappe's analysis from the Ostrom Symposium (which isn't a bad choice).
Protoavis is not discussed, nor are Protarchaeopteryx, Sapeornis or

Galton and Upchurch
23 taxa, 137 chars, CI .541
|  |--Blikanasaurus
|  `--+--Kotasaurus
|     `--+--Vulcanodon
|        `--+--Shunosaurus
|           `--Barapasaurus
      |  |--Anchisauridae
      |  |  |--Anchisaurus
      |  |  `--Ammosaurus
      |  `--Melanorosauridae
      |     |--Riojasaurus
      |     `--+--Melanorosaurus
      |        |  |(M. readi)
      |        |  `(M. thabanensis)
      |        `--+--Camelotia
      |           `--Lessemsaurus
                     |  `--"Gyposaurus" sinensis
                              |(P. engelhardti)
                              `(P. longiceps)
Ornithodira incertae sedis?- Azendohsaurus
nd- Aetonyx, Agrosaurus, Anchisaurus solus, Aristosaurus, Dimodosaurus,
Dromicosaurus, Eucnemesaurus, Euskelosaurus africanus, Fulengia,
Gigantoscelus, Gresslyosaurus, Gresslyosaurus robustus, Gresslyosaurus
torgeri, Gryponyx, Gryponyx taylori, Gryponyx transvaalenss, Gyposaurus,
Hortalotarsus, Leptospondylus, Massospondylus browni, Massospondylus
harriesi, Massospondylus hslopi, Massospondylus schwarzi, Orinosaurus,
Pachysauriscus, Pachysaurus gianteus, Pachysaurus magnus, Pachysaurus
reiningeri, Pachysaurus wetzelianus, Pachyspondylus orpenii, Paleosaurus
fraserianus?, Plateosaurus bavaricus, Plateosauravus, Plateosaurus
erlenbergiensis, Plateosaurus ornatus?, Plateosaurus plieningeri,
Plateosaurus quenstedti, Plateosaurus stormbergensis, Sellosaurus fraasi, St
renusaurus, Tawasaurus, Teratosaurus minor, Teratosaurus trossingensis,
Thecodontosaurus dubius,
Thecodontosaurus minor
Note the trend to not synonymize poorly known taxa, which is good in the
sense that a few of these 'nomina dubia' are valid.  he topology actually
isn't that disimilar from Yates', if one makes anchisaurians stem sauropods
and switches Saturnalia with Thecodontosaurus.  But the results are
questionable, given chimerical OTU's for Euskelosaurus and Sellosaurus.

Upchurch et al.
41 taxa, 309 chars, CI .514
         |  `--Neotheropoda
               |(*"Morosaurus" agilis)
                        |  |--Barapasaurus
                        |  `--+--Cetiosaurus
                        |     `--Patagosaurus
                           |  |  |(O. junghsiensis)
                           |  |  `(O. tianfuensis)
                           |  `--Tehuelchesaurus
                              |  |(M. constructus)
                              |  |(M. hochuanensis)
                              |  |(M? sinocanadorum)
                              |  |(M. anyuensis)
                              |  |(M. youngi)
                              |  |(M. jingyanensis
                              |  `(M. fuxiensis)
                                       |  |(*Amphicoelias)
                                       |  |(*Cetiosauriscus)
                                       |  |(*Dinheirosaurus)
                                       |  |(*Losillasaurus)
                                       |  |*Nigersaurus
                                       |  |(*"Cetiosaurus" glymptonensis)
                                       |  |--Nemegtosauridae
                                       |  |  |--Nemegtosaurus
                                       |  |  `--Quaesitosaurus
                                       |  `--+--Rebbachisauridae
                                       |     |  |(Rebbachisaurus)
                                       |     |  `--Rayososaurus
                                       |     |     |(R. agrioensis)
                                       |     |     `(R. tessonei)
                                       |     `--+--Dicraeosauridae
                                       |        |  |--Dicraeosaurus
                                       |        |  |  |(D. hansemanni)
                                       |        |  |  `(D. sattleri)
                                       |        |  `--Amargasaurus
                                       |        `--Diplodocidae
                                       |           |(*Dyslocosaurus)
                                       |           |(*Seismosaurus)
                                       |           |(*Supersaurus)
                                       |           |(*Tornieria)
                                       |           |--Apatosaurus
                                       |           |  |(A. ajax)
                                       |           |  |(A. excelsus)
                                       |           |  `(A. louisae)
                                       |           `--+--Diplodocus
                                       |              |  |(D. longus)
                                       |              |  |(D. carnegii)
                                       |              |  |(D. hayi)
                                       |              |  `(D. lacustris)
                                       |              `--Barosaurus
|(?*"Apatosaurus" minimus)
|  `--+--Atlasaurus
|     `--Jobaria
   |  |(C. supremus)
   |  |(C. grandis)
   |  |(C. lentus)
   |  `(C. lewisi)
      |  |(H. priscus)
      |  `(H. delfsi)
         |(*Pelurocoelus nanus)
         |(*Pleurocoelus altus)
         |*unnamed titanosauriform of Langston, 1974
         |  |--Brachiosaurus
         |  |  |(B. altithorax)
         |  |  `(B. brancai)
         |  |(?Dystylosaurus)
         |  |(Sauroposiedon)
         |  |("Cetiosaurus" humerocristatus)
         |  |("Brachiosaurus" atalaiensis)
         |  |("Brachiosaurus" nougaredi)
         |  `--Cedarosaurus
                  |(*Titanosaurus indicus)
                  |(*Titanosaurus blanfordi)
                     `--+--"Titanosaurus" colberti
                              |  `--Pellegrinisaurus
                                    |(S. loricatus)
                                    `(S. robustus)
nd- Acanthopholis platypus (in part), Aepysaurus, Algoasaurus,
Antarctosaurus brasiliensis, Antarctosaurus giganteus, Antarctosaurus
jaxartensis, Asiatosaurus kwangshiensis, Asiatosaurus, Astrodon,
Atlantosaurus, Bothriospondylus elongatus, Bothriospondylus magnus,
Bothriospondylus madagascariensis, Bothriospondylus robustus,
Bothriospondylus, Bruhathkayosaurus, Campylodoniscus, Cetiosaurus brevis,
Cetiosaurus longus, Cetiosaurus medius, Cetiosaurus mogrebiensis,
Chiayusaurus asianensis, Chiayusaurus, "Chinshakiangosaurus",
Chondrosteosaurus, Clasmodosaurus, Dinodocus, Eucamerotus, Gigantosaurus,
Histriasaurus, Hypselosaurus, Iguanodon praecursor, Ischyrosaurus,
Iuticosaurus, Kunmingosaurus, Macrurosaurus, Mongolosaurus, Morinosaurus,
Nemegtosaurus pachi,
Neosodon, Omeisaurus changshouensis, Omeisaurus fuxiensis, Omeisaurus
luoquanensis, Ornithopsis eucamerotus, Ornithopsis greppini, Ornithopsis
leedsi, Pelorosaurus becklesii, Pleurocoelus valdensis, Protognathosaurus,
Pukyongosaurus, Qinlingosaurus, Sanpasaurus (in part), Sonorasaurus,
Tienshanosaurus, Titanosaurus falloti, Titanosaurus lydekkeri, Titanosaurus
madagascariensis, Titanosaurus rahiolensis, Ultrasaurus, Zizhongosaurus
I broke from my format a bit here, as the taxa listed in parentheses
(without asterisks) under Sauropoda and Titanosauria could also go another
node up, but they are not eusauropods or lithostrotians, respectively.
Although Nigersaurus had an uncertain position in Diplodocoidea in the
analysis, the authors placed it in Rebbachisauridae in the chapter.

Ornithischians next.

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