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Heyuannia and Lu's monograph
Oviraptorid dinosaurs from Southern China
Geological Publishing House, Beijing.
2005. ISBN 7-116-04368-3. 200 pages + 8 plates.
(In chinese: pp 1-83, including 36 figures & 3 tables.
In English: pp 85-200, including 5 geological
All of the osteological figures are in the chinese
Table of contents:
Chinese section including bibliography
pp 75-83 Bibliography
pp 87 preface by Louis L. Jacobs
93 Geology of Heyuannia huangi
100 Osteology of Heyuannia huangi
100 Systematic Paleontology
112 Comparison and Discussion
116 Two new Oviraptorids from Nanxiong Basin
116 The Oviraptorid Material from Luyuan
of Shixing County
119 An Oviraptorid Dinosaur from
Xincheng Area of Nanxiong City
122 Phylogenetic Analysis of Oviraptorosauria
122 Materials and Methods
123 The Phylogenetic Analysis of
Oviraptorosaurs Based on the
Theropod Working group Matrix
130 The Phylogenetic Analysis of
Oviraptorosaurs based on Modified
143 The Status of Oviraptorosauria Among
144 Geographic and Stratigraphic Record of the
147 Summary of Conclusion
151 Appendix A - for the first phylogenetic
analysis. Character listes & character
matices for each taxon
177 Appendix B - - for the second phylogenetic
analysis. Character listes & character
matices for each taxon
199 Plate Explanation
Heyuannia huangi Lu, 2003 from the Late Cretaceous of
Southern China is
described in detail. The revised diagnosis of
Heyuannia huangi includes the following characters:
quadratojugal articular surface of the quadrate
groove-lke, the quadrate diverticulum entering the
anterolaterally, metacarpal I wrapped around
metacarpal II proximally, pneumatic
foramina present on the neural arches and ribs of
cervical vertebrae, the angle between the scapula and
coracoid approximately 145º, the ratio of
coracoid length to scapular length approximately 0.35,
the pubis as long as ischium, and the ratio of femur
length to tibia length 0.8. Two new oviraptorids from
the nanxiong Basin are also described. A phylogenetic
carried out in two parts. The first phylogenetic
analysis is to test the systematic position of
Oviraptorosauria among ceolurosaurid dinosaurs. This
analysis shows that Oviraptorosauria form a
monophyletic group. Incisivosaurus
gauthieri is the most basal oviraptorosaur and
Caudipteryx is the second most primitive form.
Alvarezsauridae is a sister group to TO
(Therizinosauroidea and oviraptorosauria). Within this
more inclusive analysis (among Ceolurosauria), all the
derived forms of Oviraptorosauria (excluding the
Primitive forms such as Incisivosaurus gauthieri and
Caudipteryx zoui) form an unresolved polytomy. The
second analysis is to determine the
interrelation-ships among oviraptorosaurs. This less
inclusive analysis (only a few groups of Ceolurosauria
were employed for analysis) allows better resolution
within Oviraptorosauria. The analysis shows
that all oviraptorosaurs form a monophyletic group,
Incisivosaurus gauthieri is the most basal
oviraptotosaur, Caudipteryx zoui is the second most
primitive form and Avimimus portentosus is basal to
the derived oviraptorosaurs. The two new taxa from
Nanxiong Basin are not nested within same clade.
Specimen BPV-112 may be the first caenagnathid
oviraptorosaur found from
Lu describes the osteology of Heyuannia in detail.
The description is based on the type specimen (the
most complete specimen) and at least 3 other
individuals. The incomplete skull was most likely
crestless (like Ingenia, Conchoraptor & Khaan). The
lower jaw is similar to other oviraptorids (being
short & deep). Heyuannia has more cervicals & sacrals
than most other small non-avain theropods. There are
13 cervicals in Heyuannia (also Richenia). Ingenia has
11 while Caudipteryx has 10. There are 8 sacrals
(Negmetia & Ingenia have 8, Nominga 5 and Richenia 7).
There are 3 sternal ribs in Heyuannia. This may be a
derived character or a result of non-preservation in
other theropods (Dromaeosauridae & Aves have 3+).
The manus is at 60 degrees to the forearm. The first
digit is very strong in Heyuannia - similar to
Ingenia. The dorsal margin of the ilium is straight
(as in Ingenia & Nemegtia). The tibia/femur ratio is
similar to many small theropods & flightless birds.
The distal end of the fibula reaches the tarsals,
unlike in Ingenia. The ascending process (of the
astragulus) is thin & tapers up.
Lu mentions two other oviraptorosaurs.
The first (BPV-112) is based on a nearly complete
sacrum, 3 posterior dorsals, 3 articulated anterior
caudals, 2 ilia and portions of the hindlimbs.
Lu refers it to Caenagnathidae gen et sp indet.
BPV-112 is a new taxon, but isn't named because of the
scarce materials. Phylogenetically, it sits closest to
Chirostenotes in Lu's analyses.
Two autopomorphies include the higher ratio of the
height/length of the ilium (not so convincing
considering the variability of ratios in specimens of
Ingenia) and the presense of a large foramen on the
proximal and of the femur.
I think BPV-112 maybe is the first definite
caenagnathid from China. It's from the Maastrichtian
The second unnamed oviraptorid is unnumbered. It is
specimen E-1 at the Nanxiang Musuem, so Lu temporarily
calls it NXMV.
NXMV is the rear half of a skull & nearly complete
lower jaw, 2 cervicals, 9 dorsals & an incomplete
right pes. Originally thought to be from Paleocene
deposits, a Late Cret age is shown from K-Ar dating
(Pingling Member of the Shanghu Formation).
So NXMV may prove to be the youngest known member of
the Oviraptoridae. Lu also decides not to name it
because it's still in preparation. Expect a more
detailed publication with a name for NXMV from Lu in
The lower jaw seems closer to Oviraptor
(philoceratops) in its proportions. The vertebrae have
well developed pleurocoels. NXMV is nested closest to
Heyuannia & Nemegtia in Lu's analysis.
Lu conducted 2 analyses. The first to test the
position of Oviraptorosauria among other theropods
including birds (using Hwang et al's ''Theropod
working group'' matrix). The second was to determine
the interelationships of oviraptorosaurs (using
Maryanska et al's matrix)
Characters used to diagnose Caudipteryx dongi seem to
be related to ontogeny, so Lu sinks C. dongi into C.
The most interesting outcome of Lu's strict concensus
tree is that Alvarezsauridae form the sister taxon to
[Therizinosauroidea + Oviraptorosauria] - far away
from a relatively 'basal' Ornithomimosauria.
>From fig 18:
Allosaurus (Sinraptor + (Tyrannosauroidea +
(Ornithomimosauria + (Ornitholestes + (Compsognathidae
+ ([Aves + Deinonychusauria] + [Alvarezsauridae +
The second analysis found the following
1)= a monophyletic Oviraptorosauria (Eighteen
synapomorphies). Includes Incisivosaurus and other (2)
taxa more derived than Incisivosaurus
2)= Caudipteryx and other (3) taxa more derived than
Caudipteryx. (eight synapomorphies)
3)= Microvenator + (4). Five synapomorphies
4)= Avimimus + (5). Three synapomorphies
5)= Caenagnathoidea. (Oviraptoroidea of Barsbold 1976.
Caenagnathoidea (Sternberg, 1940) has priority.
Caenagnathoidea is Oviraptoridae (6) (Barsbold, 1976)
+ Caenagnathidae (Sternberg, 1940)
Caenagnathidae includes BPV-112 and Chirostenotes, and
6)=Oviraptoridae includes (7)[Ingeniinae,
Oviraptorinae] & Khaan
7)= Ingeniinae + Oviraptorinae, excludes Khaan
Ingeniinae includes Heyuannia, Nemegtia, Citipati,
Richenia, Nomingia & NXMV
Within Ingeniinae, Nomingia + Richenia form a derived
group with Heyuannia, Nemegtia, NXMV, Citipati,
Ingenia forming successive outgroups to the derived
[Nomingia + Richenia] in ascending order.
Oviraptorinae includes Conchoraptor, Oviraptor & GIN
10042 (GIN 10042 has been previously discussed on the
Within Oviraptorinae, Lu weakly finds GIN 10042
closest to Oviraptor, with Conchoraptor the outgroup.
Barsbold. 1976 On a new Late Cretaceous family of
small theropods (Oviraptoridae fam. n.) of Mongolia.
(in Russian). Doklady Akademii Nauk SSSR 226: 685-688.
Hwang, Norell, Ji, and Gao. 2004. A large
compsognathid from the Early Cretaceous Yixian
Formation of China. Jounal of Systematic Palaeontology
Maryanska, Osmolska, and Wolsan. 2002. Avialan status
of Oviraptorosauria. Acta Pal. Pol 47: 97-116
Sternberg. 1940. A toothless bird from the Cretaceous
of Alberta. Journal of Pal. 14: 81-85
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