A new choristodere (Reptilia: Diapsida) from the Lower Cretaceous of western Liaoning Province, China, and phylogenetic relationships of Monjurosuchidae
Author
Gao, Ke-Qin
Author
Fox, Richard C.
text
Zoological Journal of the Linnean Society
2005
2005-11-30
145
3
427
444
https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2005.00191.x
journal article
3248
10.1111/j.1096-3642.2005.00191.x
48a5e204-fcd9-4d2b-ab85-b3cd77f813dd
0024-4082
4687363
Order
Choristodera
For more than a century since
Cope’s (1876)
naming of the group, the order
Choristodera
has been known mostly from the
Champsosauridae
and
Simoedosauridae
. The contents of the
Choristodera
, however, have been significantly expanded recently by recognition of the membership of
Cteniogenys
and
Monjurosuchus
in this group (
Evans, 1989
,
1990
;
Gao
et al
., 2000
), and by the discovery of new morphotypes including the long-necked
Hyphalosaurus
from
China
(
Gao
et al
., 1999
) and
Shokawa
from
Japan
(
Evans & Manabe, 1999
). By current understanding, the order
Choristodera
includes five families:
Cteniogenidae
,
Monjurosuchidae
,
Hyphalosauridae
,
Simoedosauridae
and
Champsosauridae
. The relationships of these families are depicted in cladograms resulting from this study (
Fig. 6
).
The taxonomically problematic taxon
Lazarussuchus
is excluded from the order
Choristodera
in this paper (see
Gao & Fox, 1998
, for discussion). Independent phylogenetic analyses consistently place
Lazarussuchus
outside the choristoderan clade (
Gao & Fox, 1998
;
Evans & Manabe, 1999
;
Ksepka
et al
., 2005
; this paper; but see also
Evans & Klembara, 2005
, for different interpretations).
Lazarussuchus
shares with choristoderes several character states including: parietal foramen absent (4-1; also absent in many other aquatic reptiles); caudal zygapophyses lie nearly or completely vertical (58-1); and fibula narrow at proximal end and flared at distal end (70-1). None of these are individually diagnosable for the
Choristodera
, and possible homologies of these characters in
Lazarussuchus
with choristoderes need to be further investigated. According to
Hecht’s (1992)
description,
Lazarussuchus
displays several features that are not otherwise known in choristoderes: paired narial openings located far back from the tip of the snout; prefrontals entering narial opening; vertebral centrum amphicoelous and notochordal; and presence of four sacral vertebrae with fused ribs. In addition, uniquely diagnostic character states, such as presence/absence of a neomorph bone in the braincase and presence/ absence of a pterygoquadrate foramen, are unknown for
Lazarussuchus
. Until these anatomical ambiguities can be clarified,
Lazarussuchus
is best treated as a taxon outside the
Choristodera
.
With the exclusion of
Lazarussuchus
, the order
Choristodera
can be unambiguously diagnosed by the following derived character states: prefrontals having median sutural contact for their entire length (2-2); parietal foramen absent (4-1; possibly homoplastic in
Lazarussuchus
and in many other aquatic reptiles); choana retracted close to midpoint of marginal tooth row (20-1); pterygoid flange consisting of pterygoid and ectopterygoid with a horizontal overlap (23-1); basipterygoid/pterygoid joint a sutural contact (24-1); parasphenoid and pterygoid having a clear sutural contact (25-1); postorbital process of jugal much shorter than anteroventral process (28-1); interpterygoid vacuity enclosed anteriorly by pterygoids and posteriorly by parasphenoid (33-1; further modification in
Champsosauridae
); neomorph in braincase present as part of external wall of braincase and medial wall of temporal fossa (40-1; unknown for
Lazarussuchus
); free odontoid process unfused to axis (50-1); vertebral centra amphiplatyan with notochordal canal closed (52-1); sacral vertebrae three in number (56-1); sacral and caudal ribs remain free from vertebrae (57-1).