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The choristoderan reptile Monjurosuchus from

the Early Cretaceous of Japan
RYOKO MATSUMOTO, SUSAN E. EVANS, and MAKOTO MANABE

Matsumoto, R., Evans, S.E., and Manabe, M. 2007. The choristoderan reptile Monjurosuchus from the Early Cretaceous
of Japan. Acta Palaeontologica Polonica 52 (2): 329–350.

The choristoderan reptile Monjurosuchus is described from the Lower Cretaceous Tetori Group of Japan on the basis of
an associated specimen from the Kuwajima Formation, Ishikawa Prefecture, and more fragmentary remains from the con−
temporaneous Okurodani Formation, Gifu Prefecture. This is the first report of Monjurosuchus from Japan, but a
long−necked choristodere, Shokawa, has already been recorded from these deposits. Monjurosuchus was first described
from the Lower Cretaceous Jehol Biota of China, although it has only recently been recognised as a choristodere. As re−
constructed, the Japanese Monjurosuchus differs from the type species, Monjurosuchus splendens, in the structure of the
postorbital region, reduction of the quadratojugal, a slender parietal with a deep groove along the interparietal suture, and
elongation of the jugal. As in M. splendens, the lower temporal fenestrae are closed. A cladistic analysis was performed in
order to place Japanese and Chinese taxa, including the incompletely described Chinese long−necked Hyphalosaurus
lingyanensis, into choristoderan phylogeny. The results support the monophyly of Neochoristodera and of a Sino−Japa−
nese clade of long necked choristoderes. The placement of the European Tertiary Lazarussuchus remains problematic,
but the analysis supports its placement within Choristodera rather than on the stem. The identification of Monjurosuchus
from Japan provides an additional link between the fossil assemblages of the Tetori Group and those of the slightly youn−
ger Jehol Biota of China.

Key wo r d s: Reptilia, Choristodera, Monjurosuchus, Cretaceous, Tetori Group, Kuwajima Formation, Japan.

Ryoko Matsumoto [ucgarym@ucl.ac.uk] and Susan E. Evans [ucgasue@ucl.ac.uk], Department of Anatomy & Develop−
mental Biology, University College London, Gower Street, London WC1E 6BT, UK;
Makoto Manabe [manabe@kahaku.go.jp], Department of Geology, National Science Museum, 3−23−1 Hyakunin−cho,
Shinjuku−ku, Tokyo 169−0073 Japan.

Introduction (Gao et al. 1999, 2000). Hyphalosaurus has 19–26 cervical
vertebrae and a short, pointed snout. Reinterpretation of the
Japan has a number of terrestrial deposits yielding fossils of enigmatic Monjurosuchus (Endo 1940) added a second cho−
Early Cretaceous age (Matsukawa and Obata 1994). Among ristodere to the Jehol Biota (Gao et al. 2000).
these, the rocks of the Tetori group of central Honshu are par− Monjurosuchus is characterized by a relatively short neck
ticularly important and have produced many well−preserved (cervical vertebral count of eight, not the usual nine) and a
vertebrate fossils. Within the Tetori Group, the Berriasian– rounded snout. Shokawa, Hyphalosaurus, and Monjurosu−
Hauterivian Okurodani Formation (Shokawa, Gifu Prefec− chus are rather unusual for choristoderes, but two more de−
ture) is the lateral equivalent of the Kuwajima Formation rived taxa, Ikechosaurus and Tchoiria, have also been re−
(Kuwajima, Ishikawa Prefecture; Maeda 1961). Both forma− ported from Lower Cretaceous formations in China and
tions are known for microvertebrate assemblages that repre− Mongolia (Efimov 1975, 1979; Sigogneau−Russell and Efi−
sent a diverse terrestrial/non−marine aquatic fauna, including mov 1984; Brinkman and Dong 1993; Efimov and Storrs
fishes, anurans, turtles, dinosaurs, pterosaurs, lizards, birds, 2000; Ksepka et al. 2005). These genera show typical choris−
tritylodontid synapsids (Kuwajima Formation only), and toderan characteristics, such as an elongated snout and rela−
mammals (Hasegawa et al. 1995; Evans et al. 1998, 2006; tively small orbits, like those of the Late Cretaceous–Early
Cook et al. 1998; Manabe et al. 2000; Barrett et al. 2002; Isaji Paleogene Euramerican genera Simoedosaurus and Cham−
et al. 2005). The choristodere Shokawa, described from the psosaurus (Cope 1876; Brown 1905; Erickson 1972, 1985;
Okurodani Formation (Evans and Manabe 1999), differed Sigogneau−Russell and Russell 1978). Most recently (Gao
significantly from any choristodere then known in having a and Fox 2005), another new small choristodere, Philydro−
long neck with at least 16 cervical vertebrae. However, an− saurus, has been described from the Aptian age Jiufotang
other long−necked choristodere, Hyphalosaurus, was subse− Formation (Jehol Group) of China. Early Cretaceous Asian
quently reported from the slightly younger horizons of the choristoderes thus demonstrate an unexpected degree of di−
Jehol Group (Yixian and Jiufotang formations) in China versity.

Acta Palaeontol. Pol. 52 (2): 329–350, 2007 http://app.pan.pl/acta52/app52−329.pdf

330 ACTA PALAEONTOLOGICA POLONICA 52 (2), 2007

N

TOYAMA

Koarashi valley

r
ive
ri R
ISHIKAWA Oarashi valley

o
Ted
“Kaseki-Kabe”

Mt Hakusan

Shokawa “Kaseki-Kabe”
Kuwajima
FUKUI GIFU 0 1 km
20 km

Fig. 1. Map of north−central Honshu, Japan, showing the “Kaseki−Kabe” locality (Kuwajima Formation) and the Shokawa locality (Okurodani Formation).
The hatched area indicates the distribution of the Tetori Group (after Maeda 1961).

New material of Monjurosuchus has now been recovered saurus, Lazarussuchus, Monjurosuchus, and Shokawa (Ev−
from the Japanese Kuwajima and Okurodani formations, ans and Hecht 1993; Evans and Klembara 2005; Gao and
Tetori Group (Matsumoto 2005). This material consists of Fox 2005). The recovery and further study of small chori−
one association and many disarticulated elements, but they stoderes such as Monjurosuchus are therefore important to
are well preserved without severe deformation. The speci− an understanding of choristoderan evolution. The purpose of
mens reveal new anatomical information for Monjurosuchus this paper is to describe new Monjurosuchus material from
and provide the first record of the genus in Japan. This is the the Lower Cretaceous of Japan, and to reassess phylogenetic
first tetrapod genus to be recorded from both the Jehol and relationships amongst basal choristoderan taxa.
Tetori groups, although the fish Sinamia (Stensio 1935) is
also known from both (Yabumoto 2005). Institutional abbreviations.—DR, Dalian Natural History
The phylogenetic position of Choristodera within Dia− Museum, China; GMV, National Geological Museum of
psida has been difficult to resolve because of the specialized China, Beijing, China; IBEF VP, Izumi Board of Education,
skull features, including a long snout, a posteriorly expanded Fukui Prefecture, Japan; IVPP, Institute of Vertebrate Pale−
skull roof, and a dorsoventrally flattened skull. Romer (1956) ontology and Paleoanthropology, Beijing, China; NMM, Na−
classified Choristodera within the paraphyletic diapsid stem tional Museum of Manchoukuo, China; SBEG, Shokawa
group Eosuchia, but most recent works have placed Choris− Board of Education, Gifu Prefecture, Japan; SBEI, Shira−
todera within the Archosauromorpha (Currie 1981; Erickson mine Board of Education, Ishikawa Prefecture, Japan.
1985; Evans 1990; Jalil 1997) or as the sister group of Other abbreviations.—CI, Consistency Index; MPT, Maxi−
Neodiapsida (Gao and Fox 1998). Within Choristodera, the mum Parsimony Tree; OTU, Operational Taxonomic Unit;
late Oligocene–early Miocene Lazarussuchus is particularly RC, Rescaled Consistency Index.
problematic. Several analyses have placed it at the base of
Choristodera (e.g., Hecht 1992; Evans and Hecht 1993; Ev−
ans and Klembara 2005), although these workers also sug−
gested that Lazarussuchus might be a derived taxon with Geology and materials
some size−linked character reversals (Evans and Hecht 1993;
Evans and Klembara 2005). On the other hand, Gao and Fox Deposits of the Tetori Group are distributed in central Japan
(1998, 2005) excluded Lazarussuchus from Choristodera on (Fukui, Gifu, Ishikawa, and Toyama prefectures; Fig. 1). They
the basis that it lacked key choristoderan synapomorphies. are composed of alternating beds of sandstone, mudstone, and
These debates are due in part to the incomplete fossil re− conglomerates of Middle Jurassic–Early Cretaceous age, and
cord of small choristoderes such as Cteniogenys, Hyphalo− show a characteristic transition from marine to freshwater

Tanankou. 2005). Kuwajima Formation is supplemented by a further specimen the Upper Jurassic–Lower Cretaceous Itoshiro Subgroup from the Okurodani Formation at Shokawa (SBEG 045). ation of right squamosal. China. are distributed in Kuwajima with only the claws projecting. diverse fossil Diapsida Osborn. complete characterized mainly (but not exclusively) by terrestrial adult individual. Matsukawa and Obata 1994). Wang and Zhou 2003). and fore− and hind feet webbed. Matsumoto et al. Lower Cretaceous (Hauterivian–Barremian. as between 140–120 Ma based on fission−track and K−Ar dat− Emended diagnosis.pdf . Niuyingzi. 1903 faunas are known from the Kuwajima Formation (Kuwajima Choristodera Cope. dark greenish−grey mudstones (Isaji et al. western Liaoning. DR0003C. 13866.—IVPP V13273. Fur− the combination of paired nasals that intervene posteriorly be− thermore. At the “Kaseki−Kabe” locality.—Choristoderan genus characterised by ing (Gifu−Ken Dinosaur Research Committee 1993). postorbitofrontal and Kuwajima formations to Berriasian–Hauterivian. 1942). 1884 [formerly Shiramine]. District. 1. 136°38’E. whereas facies III is orientalis by Endo and Shikama. Facies II commonly preserves aquatic with skin impressions (originally attributed to Rhynchosaurus vertebrates. Bessandani formations (Akaiwa Subgroup) has been dated Yixian Formation Lower Cretaceous (Hauterivian–Barremian). an adult individual (Gao et al. closed lower temporal fenestrae differentiates Monjurosuchus Fig. and fine−grained sandstone with fossil plants.—SBEI 1223 (dorsal vertebra). laterally expanded. and lizards. This squamosal). sub−adult individual over a long distance. and Japan and China. nov. China. Systematic palaeontology In the upper part of the Itoshiro Subgroup. fused frontals. the Kitadani Formation (Akaiwa Subgroup) is tween anterior tips of prefrontals. Monjurosuchus sp. group.pl/acta52/app52−329. The 1496. evidence of abrasion. vertebrates. mammals.pan. lachrymal 1977. 1940 fecture). mens are disarticulated. Several bone−bearing beds lie in the upper part of the formation and a diverse assemblage of vertebrate fossils Monjurosuchus splendens Endo. although some articulated skele− tons occur in facies III (Isaji et al. fused. and SBEI 2230 (left squamosal). Umetsu and Mat− facet on prefrontal longer than maxillary facet. and “Kaseki−Kabe” along the Tedori River (36°12’N. http://app. facies I represents a peat marsh. pls. represent three facies: facies I. Maeda 1961. the Okurodani Formation (Shokawa. a road tunnel was constructed behind Figs. dark grey fine−grained silty sandstones. Gifu Pre− Monjurosuchus Endo. the “Kaseki−Kabe”. postero− considered as late Hauterivian to Aptian on the basis of laterally expanded but markedly narrow and constricted be− non−marine molluscs and pollen assemblages (Matsukawa tween orbits. such as tritylodontid synapsids. the Kuwa− fenestra closed mainly by the enlarged jugal. 1–3. and facies Holotype: NMM 3671 (reportedly lost during World War II). Hakusan City (formerly Shiramine Village). 1940 Monjurosuchus splendens sp. It is impossible to give a precise orig− Tetori Group is divided into three subgroups: the Middle–Up− inal location for these specimens.. medium− al. and 2230). facies II a shallow lake. 2005). Fossils show little Referred specimens. nov. suggesting they have not travelled small juvenile individuals. 13279. vertebrates. 2000). Isaji 1993. a short rounded preorbital region. 2–15. western Liaoning. SBEI 1496 (left were from the upper part of the Kuwajima Formation. 2. The Kuwajima Formation is overlain conformably from all other described choristoderes (modified from Gao et by the Akaiwa Formation and consists of massive. The rocks drilled out during construction Material. frontals fused. Lower Cretaceous (Hauterivian– preserves a variety of vertebrate remains and most speci− Barremian) Yixian Formation (Zhou et al. 2000). 14269. SBEI 1792 (block bearing an association of material was preserved and set aside for study and includes bones). grain size and the fossils (plants and molluscs) indicate that Yixian Formation. pls. Each facies Neotype: GMV216. Lingyuan. 2005). Here they form a large bluff known as the only eight cervicals. facies II. and facies III a vegetated swamp (Isaji et al. supratemporal suoka 2003). prefrontal equal to frontal in length. The approximately 10. III. Ishi− The combination of narrow. 1940 have been collected from the “Kaseki−Kabe” locality. infratemporal The uppermost part of the Itoshiro Subgroup. jugal. Tanankou. (Tithonian–Hauterivian) and Akaiwa Subgroup (Barremian– Albian. 2002). such as fishes and turtles. Kusuhashi et al. 1). These data limit the age of both the Okurodani fenestra roughly equal to orbital length. and quadrate). squamosal square. carbonaceous sandstones. and the lowermost part of the Akaiwa Sub− shaped with straight stem. The material from the per Jurassic Kuzuryu Subgroup (Bathonian–Kimmeridgian). molluscs. IVPP V3673. 2003).—CRETACEOUS CHORISTODERE FROM JAPAN 331 through brackish water environments (Maeda 1961). 1982. SBEG 045 (associ− four of the specimens described in this paper (SBEI 1223. interclavicle “T” jima Formation.MATSUMOTO ET AL. China. a short neck of kawa Prefecture. Ishikawa Prefecture) and its lateral equivalent.—Lower Cretaceous (Berriasian–Barremian). the Akaiwa Formation. The section spanning the Okurodani and overlying Type species: Monjurosuchus splendens Endo. From 1997 to 2000.. to fine−grained sandstone and alternating beds of siltstone Distribution. 1792. Endo 1940: 1–14. quadratojugal. Endo and Shikama 1942: 2–6. Terrestrial and freshwater vertebrate fossils are known from all sub− groups (Azuma and Tomida 1995. 1940. the bone−bearing beds 1942 Rhynchosaurus orientalis sp.5 km southwest of Lingyuan.

these a nearly complete right surangular is associated with the an− are always left and right. Japan. It is morphologically iden− the fact that the quadrate−articular joint lies forward of the tical to the other squamosals. 3–12). are not identified in SBEI 1792. SBEG 045. prearticular. As reconstructed. Choristoderan reptile Monjurosuchus sp. With one exception (see below) some confidence. 2. as are the vertebrae. Only posterior part of the right frontal. The dermal bones are ornamented from a second individual. (SBEI 1792). However. a right quadratojugal. premaxilla. the lateral part of the right squamosal. As a re− splenial. articular.—In Choristodera. but some preorbital elements are not. It is therefore highly probable that the bones on SBEI the length of the mandible can be reconstructed as approxi− 1792 constitute the remains of a single individual. the upper surface of the a left splenial. it is not formally named herein. premaxillae. pari− surangular. mm × 140 mm × 20 mm (Fig. The presence of this second indi− with a pattern of anteroposterior striae. Many mately 70 mm. Description. surangular. and qua− postorbitofrontal. specifically distinct from the Chinese Monjurosuchus splen− right and left dentaries (associated angular). dorsal. an associated right prefrontals.—SBEI 1792 is a block of dark greenish−grey vidual renders SBEI 1792 problematic as a potential holo− mudstone preserving 25 choristoderan bones including: cer− type. quadratojugals. In SBEI 1792. Most of the bones are at 4). the drate. and the posterior part of the quadrate are preserved on SBEG 045. an articulated dens. postfrontals (or combined postorbitofron− right parietal. lachrymals. maxillae. SBEI 2230. squamosal. the partly articulated postorbital elements least partly articulated. The smaller. the lachrymal.332 ACTA PALAEONTOLOGICA POLONICA 52 (2). Photograph of main block of the holotype. Ishikawa Prefecture. an articulated left squamosal. postorbitals. but parts of the lateral part of the right frontal. 2A). As seen in dorsal . there are never more than two examples of each skull ele− However. and right dentary are iso− sult. A and B show alternate sides of the block. 3). 2). the skull preserved on other associations of this type are known from the same de− SBEI 1792 is 80 mm in length and about 50 mm wide. splendens (DR0003C) is shown for comparison (Fig. squamosals. more fragile cranial elements and The skull is reconstructed on the basis of SBEI 1792 and most of the postcranium are not preserved. jugals. and quadrates. The posit. Fig. size. and clearly came posterior limit of the skull. In addition. and angular. but smaller. The bones on isolated specimens (SBEI 1496. Kuwajima. broken and their parts attached to neighbouring bones). 2007 squamosal (SBEI 2230) squamosal (SBEI 2230) 10 mm prefrontal prefrontal dorsal vertebra mandible splenial surangular parietal prefrontal maxilla surangular caudal vertebra mandible maxilla quadratojugal mandible postorbitofrontal Fig. SBEI 1792 preserves a nearly complete right den− ment on the block (allowing for the fact that some bones are tary and a left dentary with an associated angular. the anterior part of the right squamosal. the postorbital skull length can be reconstructed with lated. left maxilla and the anterior part of a jugal. but the preorbital region is more difficult. right and left prefrontals. an articulated etals. and an articulated right four elements. frontals. Lower Cretaceous Kuwajima Formation. A skull SBEI 1792 form an association within a small area of 110 of M. nasal. Skull (Figs. and although the Japanese Monjurosuchus is probably vical. jugal and skull is composed of eleven or twelve pairs of bones: nasals. the left are preserved. and caudal vertebrae. right jugal. quadratojugal. From these elements. and the tal). and they are always of matching gular and a small part of the dentary. is difference between jaw and skull length can be explained by also found on the block (Fig. an additional left squamosal. a left surangular. but four skull elements.

The dorsal sur− face is horizontal and sculptured. China. in lateral (A). Kuwajima. and suggests that the suture between parietal the squamosal and parietal lay half way along the postero− medial border of the upper temporal fenestra. and covers http://app. Choristoderan reptile Monjurosuchus sp.pdf . The prefrontal tapers posteriorly. an anteroposterior flange is seen to bear a medially inflected notch−like facet for the lachrymal. Japan. 3A). The implanta− tion is subthecodont and tooth sockets are ovoid. and its narrow frontal process Fig. Choristoderan reptile Monjurosuchus sp. The quadrate is located behind the squamosal/post− 10 mm squamosal orbitofrontal suture so that the lower jaw is posteriorly elon− postorbitofrontal quadratojugal gated in comparison to that of Monjurosuchus splendens. The elongated right and left prefrontals are both pre− served in SBEI 1792 (Fig. 5A).MATSUMOTO ET AL. (SBEI 1792). Liaoning Province. 6A).—CRETACEOUS CHORISTODERE FROM JAPAN 333 postorbitofrontal squamosal parietal squamosal prefrontal 10 mm parietal maxilla jugal frontal lower jaw maxilla jugal postorbitofrontal quadratojugal Fig. the squamosal lacks the process for the parietal so the length of parietal process was inferred from an isolated nasal complete squamosal (SBEI 1496). 3B). dorsal (B). 6A). Ishikawa Prefecture. 5. and part of the post− orbitofrontal lie vertically and form the lateral side of the skull jugal (Fig. In dorsal view (Fig. Partial reconstruction of the skull in dorsal (A) and lateral (B) views. In prefrontal SBEI 1792. but all teeth have been lost (Fig. and partial ventral (C) views. 5). and is sensory foramina 5 mm partly lacking the frontal process (Fig. Yixian An incomplete left maxilla is preserved in SBEI 1792. 5B). the length of the upper temporal fenestra is similar to that of the orbit. with a well−developed or− bital flange bearing weak tubercles (Fig. This is limited to the anterior tip of the bone. 4.pan. quadratojugal. Ishikawa Prefecture. Circle and dash missing the regions for articulation with the premaxilla and pattern indicates matrix. As recon− structed. but the ventral side is covered by matrix (Fig. 6). The left bone is of equal size but has a crack in the middle. The maxilla is mediolaterally narrow and dorsoventrally flat− tened. Choristoderan reptile Monjurosuchus splendens (DR0003C). frontal view (Fig. Skull in dorsal view. The squamosal process on the parietal is complete. jugal. Lower Creta− enters the rim of the large. 5C). the flange articulates with the orbital ramus of maxilla sensory foramina jugal the jugal. The straight me− dial edge bears a deep suture for the opposite prefrontal. Posteriorly.pl/acta52/app52−329. Formation. dorsolaterally directed orbits. maxilla The squamosal. 6). with its lateral surface ornamented with a pattern of fine anteroposterior striae and sensory foramina (Fig. Left Anterodorsally. ceous Kuwajima Formation. 6B). 3. Kuwajima. there is a small recess for the nasals (Fig.. Fig. Lower Cretaceous Kuwajima Formation. Japan. maxilla and jugal. the orbits are large and dorsally directed and lachrymal the upper temporal fenestrae are anterolaterally expanded. At least seven tooth sockets are visible on the lachrymal facet maxilla. prefrontal but with part of the jugal in association (Fig. The right bone is nearly com− plete.

Choristoderan reptile Monjurosuchus sp. lateral (B). 2007 nasal facet maxilla facet in all known choristoderes. jugal. Laterally. It lacks the tip of ceous Kuwajima Formation. 8). ceous Kuwajima Formation. 5A. Lower Creta− parts of the frontal. Ishikawa Prefecture. Kuwajima. jugal only one fifth of the entire prefrontal length. the jugal is anteroposteriorly elongated squamosal frontal postorbitofrontal 5 mm squamosal squamosal Fig. the postorbital and postfrontal are fused without trace of a suture (Fig. Japan. (SBEI 1792). 8. in dorsal (A). 8. and is limited by the anterior part of an elongated squamosal lachrymal facet that lies parallel to the interprefrontal suture. Japan. 9). The right jugal is better pre− parietal served and is associated with the postorbitofrontal and squa− mosal. A1. parietal. A deep concavity runs parallel to lachrymal facet the midline (Fig. Right the frontal facet and the neomorph process. As in all choristoderes except Champsosaurus and Ctenio− genys. Kuwajima. Lower Creta− ceous Kuwajima Formation. and ventral (C) views. 6B). B. Choristoderan reptile Monjurosuchus sp. The main body of the bone contacts the squa− palatine facet mosal and jugal and forms the posterior margin of the orbit 5 mm and the anteroventral margin of the supratemporal fenestra (Fig. 7A). As seen in medial view. The left and right jugals are preserved in SBEI 1792 lateral process (Figs. but its orbital flange and quadratojugal facet are miss− ing (Fig. . The maxillary facet is extremely postorbitofrontal reduced. Overall.334 ACTA PALAEONTOLOGICA POLONICA 52 (2). The anterior part of the left jugal is associated with the maxilla and the posterior part is associated with the quadratojugal and squamosal. the medial edge forms the margin of a channel lead− ing forward from the frontal into the posterior nasal cavity (Fig. and squamosal. B2. 6B). A dorsally expanded wing meets the frontal and pa− rietal and contributes to the posterodorsal margin of the orbit Fig. maxilla facet ectopterygoid facet SBEI 1792 preserves the right parietal in association with Fig. but the middle section with facets for the maxilla and postorbito− frontal is missing (Figs. C). but are fragmented. Below the lachrymal facet lies the palatine process (Fig. 3. 7A). (SBEI 1792). (SBEI 1792). Lower Creta− and the anterior margin of the supratemporal fenestra (Figs. 6B). 7A. in ventral view. Anterior and lateral processes met the frontal and postorbitofrontal respectively (Fig. and is longer than the squa− mosal facet in lateral view (Fig. 8A). 8B). 6. The external surface of the postorbitofrontal is sculptured prefrontal in dorsal view. separated from the posterior nasal cavity by the medial edge of the palatine process (Fig. photographs. 8). Pos− posterior terolaterally. Right 8A). Ishikawa Prefecture. the parietals are paired and lack a anterior cavity parietal foramen (Fig. As (B) views. 7). Right frontal. and squamosal (Fig. 9A). B1. the parietal is drawn into an expanded squa− cavity mosal wing that has a strong overlapping facet with the squamosal (Fig. Kuwajima. but is otherwise frontal. A. 7. in lateral (A) and medial nearly complete. Choristoderan reptile Monjurosuchus sp. postorbitofrontal. 7A. there are anterior and posterior facets for the maxilla and lachry− 5 mm mal respectively (Fig. and the dorsolateral edges are tuberculate (Fig. frontal the anterior orbital rim is strengthened by buttresses but be− frontal hind this the bone becomes very thin (Fig. interpretive drawings. The resulting postorbitofrontal is divided into two parts. A2. Japan. and squamosal. The jugal facet is anteroventrally expanded. The bone is anteroposteriorly elongate. B). 6A). Left prefrontal. 8A). Anteropos− jugal teriorly. Ishikawa Prefecture. 8A). The anterior nasal cavity terminates in the middle of the pala− frontal tine process.

but supple− and dorsally expanded. This parietal Fig. 9A–C). D. as shown in SBEG 045 (Fig. 8B). the quadratojugal−jugal suture is a straight line. B4). and neomorph. 8. with spike−like tubercles ornamenting the dorsal and panded flange that forms the curved orbital margin (Fig. but tion of the right quadrate. with an uneven surface (Fig. an associa− gin. 8). while posteriorly it smooth surface. 10A2. On SBEI 1792. (SBEI 1792). with a narrow dorsal apex bital rim (Fig. The bone is attached to the temporal fenestra. In lateral view. 8A. B1. The squamosal is made up of lateral and medial plates. 10). D).MATSUMOTO ET AL. 9C). the smaller left bone on the same block. The medial and lateral squamosal plates contact each other vertically (Fig. and quadratojugal (Fig. 9A). The squamosal has a strongly sculptured lateral gular cross−section (Fig. quadrate facet The dorsoventrally expanded lateral plate forms the side of squamosal the skull and the lateral margin of the supratemporal fenestra facet ?rib (Figs. 10B3–B6). and it is also missing the parietal process. photographs. B). and is seen to be ceous Kuwajima Formation. 10B3–B6). Anteriorly. quadratojugal. interpretive drawings. 8). A ?rib medially expanded flange gives the bone an “L” shape in ventral view (Fig. Its dorsal and ventral margins run almost in parallel.pl/acta52/app52−329. The medial ectopterygoid facet lies above capped by a small cephalic condyle (Fig. each separated by grooves (Fig. A2. B4). Scale bars 5 mm. 9A. C. medial plate is inclined posterodorsally at 45 degrees (Fig. Drawing of right quadratojugal in lateral view.pdf . and 10A3). It lacks the dorsal margin and the parietal pro− cess. 8B). A–C. squamosal. facet running beside it (Fig. in lateral (A). This flange meets the squamosal dor− jugal quadratojugal sally and the quadrate posteriorly. SBEI 2230. 10B4). 10A) and SBEG 045 (Fig. It gives rise to a dorsally ex− surface. 9A. A2. the quadrate. Left squamosal. 9A. In the postorbital region. quadratojugal. 7) and its lateral wing associated with the jugal and postorbitofrontal (Fig. 9A–C). the squamosal facet being roughly equal quadratojugal posterolaterally. The medial plate quadratojugal jugal forms the posterior rim of the supratemporal fenestra and ex− tends to meet the parietal (Fig. and the squamosal but does not contact the postorbitofrontal. B). and pos− the ventral edge is much thicker. 10B3–B6). The The quadratojugals are preserved on SBEI 1792 (Fig. jugal. Dorsolaterally. this parietal process bears a small concavity for the quadrate and the quadratojugal. but the surrounding facets indicate that no meets the squamosal and quadratojugal to close the lower movement could have occurred. terminating just behind the level of the or− ments. although the right bone is split. The qua− the jugal laterally (Fig. ornamented with the characteristic sculpture pattern and a tuberculate ventral margin (Fig. and jugal. the squamosal dorsally.pan. 10B3–B6). medial (B).—CRETACEOUS CHORISTODERE FROM JAPAN 335 matrix (Fig. 9A. and its medial side is covered with there is a shallower surface for the opisthotic. Japan. Both squamosals are preserved on SBEI 1792 (Figs. 10B). the plate has sutures with the postorbitofrontal. The quadratojugal is reduced to a small tri− squamosal angular plate. B3. B2). and in length to the quadratojugal facet (Fig. the jugal is mediolaterally flattened The quadrate is not preserved in SBEI 1792. condyle is located near the lateral surface of the skull and has a the jugal overlies the postorbitofrontal. with the deep. although the with a strongly sculptured lateral surface (Figs. 9A–C). Kuwajima. Ishikawa Prefecture. The jugal bears a number of terior part of the jugal (Fig. The left squamosal is articulated with a jugal and a quadratojugal (Fig. The anterior process is dorsoventrally flattened with a trian− 9B. 9D). but SBEI quadratojugal 1496 (Fig. pterygoid. rather square in dorsal view. 10B5). 10B3. It artic− ulates with the jugal. the left bone is up the medial edge. B2). subequal in length to the quadratojugal−squamosal suture (Fig. Lower Creta− process is well−preserved in SBEI 1496. Its medial face is broad and drate facet is weakly striated and is located just above the bears a series of interlinked facets for the opisthotic. B). with its anteromedial process attached to the parietal (Fig. 5A. The ventromedial maxillary facet lies along SBEI 1792 in the shape and the size of the component ele− the ventral margin. A1. posteroventral pterygoid facet extends about half way and SBEG 045 (Fig. is jugal isolated. Choristoderan reptile Monjurosuchus sp. B1. B4). From anterodorsal to postero− ventral. 9) large. The quadrate is triangular. 10B3. B2. forming the vertical lateral skull mar− mentary information is available from SBEG 045. anteroventral neomorph complete and has an expanded medial surface (Fig. 9C). 10A1. forming a deep quadrate squamosal recess (Fig. 9A. http://app. ventral (C) views. 10B) are almost com− plete. 9C). B1. 9. Above these two facets the right bone is detached. junction of the jugal and quadratojugal (Fig. The the maxillary facet as a weak notch (Fig. posterior rims (Figs. SBEG 045 matches distinct facets. 10A5.

the symphysial surface is short by but four are preserved within the alveoli (Fig. B4–B6. A1. dorsal (A3). 11A1). six (the dentary. 11). Here the splenial facets end and dentary is a slender element with the anterior end slightly up− are replaced by the angular facet in the floor of the Meckelian turned. 11A2). B1. jugal. in lateral (A1.336 ACTA PALAEONTOLOGICA POLONICA 52 (2). splenial. B. The depth gradually increases posteriorly (Fig. B2). canal is not preserved. and Lazarussuchus. symphysis. and posterior parts (Fig. part of the angular is The external surface is finely striated. B3. quadratojugal. 2007 postorbitofrontal facet quadrate facet quadrate facet quadratojugal facet jugal facet postorbitofrontal facet parietal process squamosal quadrate facet jugal quadratojugal quadrate opisthotic facet quadrate quadrate jugal quadratojugal squamosal neomorph facet 5 mm pterygoid facet quadratojugal Fig. 11A2). The tooth implantation is subtheco− nerve foramina. from the symphysis. in lateral (B1. each of which opens into a short groove (Fig. dont as in all Choristodera. A2–A5. Its dorsal and ventral margins run in parallel. 11B). Of these. Japan. angular. medial (B3. leaving the anterior part of the fossa splenial. Japan. Lower Cretaceous Kuwajima Formation. B2. The Meckelian fossa is enclosed at the Hyphalosaurus. fossa (Fig. 11A2). as seen in Cteniogenys. The bone is flat and thin with This resembles the condition in the Jurassic Cteniogenys. chus (Fig. B4). articular. and surangular) have open (Fig. prearticular. and quadrate (SBEG 045). . but is lacking its anterior the subdental shelf is thicker than the lower edge (Fig. and subdental ridge and the lower border of the Meckelian fossa a partial left dentary articulated with an angular (Fig. Okurodani Formation. interpretive drawings. Ishikawa Prefecture. and ventral (B6) views. On the left dentary. suran− Meckelian fossa and show that the splenial was excluded gular.—In choristoderes the lower jaw is made up of Facets for the splenial are clearest on the lower border of seven bones: dentary. Kuwajima. Shokawa. and medial (A5) views. 11A1). A2). The narrow abruptly (Fig. and articular. Towards the rear of the tooth row. but The left splenial is preserved. coronoid. Gifu Prefecture. ventral (A4). Most of the teeth have been lost. posterior (B5). angular. Lower jaw. In medial view. Right squamosal. 11A). 11A2). with a line of sensory associated (Fig. 11A2). the SBEI 1792 preserves a nearly complete right dentary. Left squamosal (SBEI 1496). straight dorsal and ventral margins. They comparison with that of neochoristoderes and Lazarussu− are conical with a smooth base. 11C). A. prearticular. 10. and a smooth surface. The entry foramen of the inferior alveolar been identified in SBEI 1792. Choristoderan reptile Monjurosuchus sp. photo− graphs.

Ishikawa Prefecture. articular. http://app. Both surfaces are sculp− tured. 12A2) is robust and pos− dentary teriorly expanded. extending angular facet medially into the Meckelian fossa (Fig. B1). (A2) views. and Lazarussuchus. the articular fossa extends over a A. angular. lying above the posterior part of the pre− articular. views. the base of which is robust. surangular SBEI 1792 also preserves the left and right surangular. but its midsection is preserved in association with the dentary (Fig. The articular (Fig. B2. angular. Scale bars 5 mm. The surangular is a robust element that lies along the posterodorsal edge of the mandible. and the posteroventral margin is also tuberculate. in lateral (A1) and medial larger area and fills almost the entire depth of the surangular. the latter facet angles dorsally and is bordered by coronoid facet a raised. ventrolateral angular facet (Fig. B1). and dorsal (B3) There are no nerve foramina on the medial surface. posteriorly. B3). C2. Left surangular. (SBEI 1792). B. Japan. Choristoderan reptile Monjurosuchus sp. 11C2). 12A2). 13. B. 11C). Circle and dash pattern in A1 indicates matrix. In medial view. robust element that contributes to the posteroventral border of the mandible. and lateral sculpture angular (Fig. Anteriorly it meets the dentary. (SBEI 1792). Lower choristoderes.—CRETACEOUS CHORISTODERE FROM JAPAN 337 The right angular articulates with the surangular and prearticular (Fig. 12B). 12A2). tuberculate rim. occupying only the upper half of the surangular (Fig. 12A). 14). Japan. the left is more fragmentary. The bone bears a deep anterodorsal coronoid facet and a broad. and Tchoiria. Hyphalosaurus. C. with a weakly sculptured medial sur− angular face (Fig. 12A1. and prearticular (Fig. is the facet for the articular (Fig. The left bone is isolated. Two lingual sensory foramina open anteroventrally (Fig. Left dentary in coronoid facet medial view. covering the sensory foramina ventral margin of the surangular. Postcranial skeleton (Figs. 12A). The angular is a long. This resembles the condition in articular facet Cteniogenys. Lower surangular Cretaceous Kuwajima Formation. interpre− tive drawing. B3). It is therefore exposed both medially and laterally (Fig. Choristoderan reptile Monjurosuchus sp. B3). The dorsal margin bears a slight posteromedial inflec− 5 mm dentary facet angular tion. and mostly internal to both it and the surangular (Fig. A. In neo− Fig. 11. There is a narrow groove facet for the prearticular. and prearticular. Left splenial in lateral view. Simo− Cretaceous Kuwajima Formation. 12B2. Ikechosaurus. photograph. Just above it.pdf . and is associated with the articular dentary. The right bone is nearly complete. It runs above the angular and is anteroposteriorly elongate and slender.MATSUMOTO ET AL. Fig. the anterior ad− angular facet ductor chamber is separated from the posterior articular re− gion by a smoothly curved crest (Fig.pl/acta52/app52−329.—The only postcranial elements that can be attributed with confidence to the Japa− sensory foramina nese Monjurosuchus are vertebrae. Posteriorly. The right prearticular is associated with the articular (Fig. and is missing its anterior part (Fig. Right dentary in lateral (A1) and medial (A2) views. in lateral (B1). C1. The dorsal border of the adductor compart− articular facet prearticular facet ment is thick and posteriorly rounded. 12A2. 12A2). such as Champsosaurus. Kuwajima. 11B). 11B). 12. Ishikawa Prefecture. Right surangular. 12A). 12A2). 12B2). edosaurus. It is almost rectangular splenial facet prearticular with parallel dorsal and ventral margins. probably to accommodate the coronoid (Fig. 12A1.pan. Kuwajima. 12B2. The articular fossa is relatively shallow. The angular runs parallel to the prearticular and narrows posteri− splenial facet orly (Fig. whereas the ventral border is incurved (Fig. medial (B2).

mandibular foramina open into anteroposteriorly oriented grooves. This is slightly larger than the vertebrae on SBEI 1792. 14A. and margin of lower jaw anteroposteriorly orientated at 45° to the horizontal. Japan. positioned just above the level of the neuro− caudal rib central suture. ?spinous process Although two fragmentary dorsal vertebrae are preserved on SBEI 1792. interpretive drawing. in left lateral (A1). The posterior facet. C). 13A2). Fig. Choristoderan reptile Monjurosuchus sp. and platycoelous anterior vertebral centrum (Evans and Klembara 2005). suchus in having the following characters states: amphipla− with both the neural spine and anterior zygapophyses broken tyan vertebral centrum with a closed notochordal canal. limited to one or two zygapophysis is a broad. 14. with the posterior zygapophysis orientated at 45° to the horizontal plane (Fig. Lower Klembara 2005). in left lateral (A). anterior (B). flared posterior tem− poral region that extends posteriorly. Evans and Fig. The centrum is platycoelous and is excavated by a ceous Kuwajima Formation. by far the most complete specimen is SBEI 1223 (Fig. The anterior and posterior zygapophyses are roughly equal in length (Fig. Cer− shallow concavity on each side of the midline. SBEI 1792 differs from Lazarus− A single cervical vertebra is preserved on SBEI 1792. 13A2). Ishikawa Prefecture. a shorter mandibular symphysis. photograph. 2007 process of the dorsal vertebrae (Fig. (SBEI 1792). 14). This spinous rib is rectangular in cross−section and. SBEI 1792 differs from Hyphalosaurus in the follow− is a small expansion that may be equivalent to the spinous ing characters: the cervical neural arch is taller.—SBEI 1792 presents the following choristoderan synapomorphies: median contact of prefrontals along entire length of elements. flanked on either side by lateral concavities. and posterior (C) views. Lower Creta− 14A). Kuwajima. The parapophysis is shorter and is horizontal in orientation. Part of the caudal rib is fused to the centrum. 13A1). and the A1. in lateral (B1) and ventral (B2) views. in these two characters SBEI 1792 shows similarities to the Oligocene to Miocene European Lazarussuchus (Hecht 1992. Ikechosaurus. 14A). and ventral (A2) views. Japan. 13A). and Tchoiria) and the Jurassic Cteniogenys. elongated postorbitofrontal. However. saurus (Gao et al. and it bears acces− sory facets. Simoedosaurus. 13B). B. rectangular plate (Fig. 1999. The centrum has a strong midventral keel (Fig. It is longer than the cervical and dorsal ver− tebrae. 14C). zygapophysis SBEI 1792 is distinguished from all neochoristoderes (Cham− psosaurus. and Philydro− saurus (Gao and Fox 2005). parietal foramen absent. just anterior zygapophysis above the level of the neurocentral suture (Fig. The neural spine has an apex with a smooth surface. 2000. A. as preserved. (SBEI 1223). The transverse process is positioned above the centrum (Fig. Scale bars 5 mm. the Early Cretaceous Chinese Hyphalo− Cretaceous Kuwajima Formation. poste− (Fig. 13B1) and bears a midventral groove for caudal blood vessels (Fig. spinous process conical subthecodont teeth. The centrum is concave in lateral view (Fig.338 ACTA PALAEONTOLOGICA POLONICA 52 (2). There is a small spinous process be− low the postzygapophyses (Fig. Below it on the right side straight. 13B1). Its tip is vertically expanded and bears a posteriorly concave rib facet (Fig. the dorsal . Kuwajima. coarse exter− 5 mm nal sculpture. Evans and Klembara 2005. 13A1). vical vertebra. Ishikawa Prefecture. Choristoderan reptile Monjurosuchus sp. Partial caudal ver− A partial anterior caudal vertebra is preserved on SBEI tebra. 13A1). 13B 1). is long process and robust although it is broken in the middle (Fig. and is tooth positions. although this is closed without a trace (Fig. with no riorly constricted prefrontal with a much shorter maxillary trace of the neurocentral suture (Fig. slender dentary. 1792 (Fig. Early Cretaceous Chinese Monjurosuchus (Gao et al. Furthermore. Dorsal vertebra. Gao and Fox 2005). 13A2). Remarks. in the closure of the lower temporal fenestra and a posterior constriction of the prefrontal that is as− sociated with medially expanded orbits. RM personal observation). The transverse process is expanded and sin− gle−headed. 13B2). striated on maxillae. but has the same morphology. The neural arch is fused to the centrum. 13. A2.

splendens. the compound postorbitofrontal met the frontal vical vertebra (laterally expanded in Shokawa). In Cteniogenys. In SBEI 1792. 13279. 4). Posteriorly. and the dor− some characters. tures are closed (open in Shokawa with centra that are nearly In the new Japanese choristodere. and differs symphysis. and ventral margins of the surangular are subparallel. Further− However. the maxilla that is limited to the anterior region between the splendens in its shape and in the anteroposterior expansion of nasal and lachrymal facets. and the The parietal of SBEI 1792 resembles that of Cteniogenys articular flange and symphysis lie on the same horizontal (Evans 1990). vaded by the nasal to about one quarter or one fifth of its spanning the length of 1–2 tooth positions. Hyphalosaurus. the squamosal is more slender sal surface of the parietal is flatter. and IVPP (IVPP V 3673. in SBEI 1792. splendens and the Japanese choristodere differ in rietal of M. having the facets parallel to each Manabe 1999). in splendens. In Neochoristodera. the dentary is more slender. ches to the skull margin. and the frontal and frontal facets lie on the same level as the inter− prefrontal has a broader maxillary contact. the length of the quadrato− by Gao et al. Evans and Klembara 2005). and postorbitofrontal of both taxa are of similar shape. Hecht 1992. The dorsal vertebrae have verti− several differences between the parietal of SBEI 1792 and cally expanded single−headed transverse processes. splendens. dorsal. The jugal meets the quadratojugal 13866. the lower temporal equal in size). The prefrontal of SBEI 1792 matches that of anterior to the squamosal/postorbitofrontal junction in M. the This is shorter than that of Lazarussuchus (roughly six tooth process for the frontal is markedly narrow and constricted. length.pdf .pl/acta52/app52−329. splendens (RM personal observation). Fig. How− above the level of the interparietal suture. and in lacking the that the lateral part of the quadrate mandibular condyle atta− distinct supratemporal trough (Gao and Fox 2005). parietal suture. but jugal is roughly one half that of the jugal. M. Hyphalosaurus. Hyphalosaurus (RM personal observation). The latter is extremely long and the parietal process. as in SBEI 1792. 2000). 13273. no skull material is yet avail− differs from that of Cteniogenys in the position of the neo− able for the Early Cretaceous Japanese Shokawa (Evans and morph and quadratojugal. the postorbitofrontal contacts the squamosal anterior to ture (horizontally and just on the neurocentral suture in the mid−point of the supratemporal fenestra. 4). The only postcranial elements known for the Japanese matching the condition in M. there is a small spinous http://app. the parietal is longer coelous cervical. the interprefrontal suture is in− served in IVPP V13279. the dorsal late instead. posterior zygapophyses expand had a closed lower temporal fenestra (Gao et al. The size and shape. and the quadratojugal facet is lim− ton: the cervical neural arch is taller. splendens. The quadrate of the Japanese reptile Apart from attributed jaws. dorsal neurocentral su− ited to the posteroventral edge. but not in M. as in SBEI 1792. the surangular is posteroventrally Lazarussuchus (Hecht 1992. deflected and the articular flange lies below the level of the and M. but The new Japanese choristodere is very similar to Mon− the posterior part of the jugal and the quadratojugal of the jurosuchus splendens (Fig. splendens (IVPP V13279) is that of M. the prefrontal of SBEI 1792 differs slightly from more. SBEI 1792 resembles that of Cteniogenys. but not other choristoderes. In in parallel with each side. positions. the neomorph facet lies between the from Shokawa based on characters of the postcranial skele− pterygoid and opisthotic. Monjurosuchus splendens also flank grooves in Shokawa). However. However. SBEI 1792 can be distinguished other. splendens in based partly on the description of Monjurosuchus splendens shape and length. unlike that of Cte− anterolateral expansion that forms most of the skull roof. splendens. Monjurosuchus splendens. At the an− The mandible of Monjurosuchus splendens is partially pre− terior tip of the prefrontal. as seen in a dorsal view of the cer− both taxa. extends along the lateral margin of the prefrontal. 14269). splendens is anteroposteriorly short. dorsoventrally expanded. in having suchus splendens differ from those of neochoristoderans in distinct ornamentation on the squamosal. so that the postorbito− in its posterior part. and Lazarussuchus. In M. the pa− ever. Evans and Klembara 2005). In M. the ventral margin of the from Philydrosaurus (Gao and Fox 2005) in having an surangular is robust and curves medially. The niogenys.pan. and caudal centra with fully closed anteroposteriorly and more slender. The surangular of which bears a steep dorsally directed flange. The nasal facets are wedge−shaped. centra of caudal vertebra bear a shallow ven− fenestra is closed by the elongated jugal and expanded post− tral groove for caudal blood vessels (deep ventral flanges orbitofrontal and squamosal. In contrast. and platy− that of M. The following comparison is Japanese choristodere differ from those of M. In the Japanese form. In lateral diapophysis orientated vertically and above neurocentral su− view.MATSUMOTO ET AL. This combination and frontal facets are borne on a dorsolateral flange that lies of characters matches only Monjurosuchus splendens. the posterior dentary of M.—CRETACEOUS CHORISTODERE FROM JAPAN 339 neurocentral sutures are closed. The squamosal Shokawa). splendens in the structure of the orbital margin. (2000) and Evans and Klembara (2005). and cervical anteromedially and the parietal posteromedially. SBEI 1792 differs from the new Philydrosaurus in lack− The quadrates of the Japanese choristodere and Monjuro− ing distinct ridges on the prefrontal and postfrontal. the orbital margin lacks this flange and is strongly tubercu− and Lazarussuchus in general shape. splendens. there are choristodere are vertebrae. whereas an elon− mainly on a study of new complete material in DR gated jugal meets the squamosal in the Japanese choristodere (DR0003C. Both share a prefrontal with a single facet for squamosal of the Japanese choristodere matches that of M. and preserves a terminal symphysis. but behind it in the Japanese choristodere. parietal of SBEI 1792 has a broad facet for the squamosal. In these taxa. However. plane. and the postorbitofrontal neurocentral sutures and no notochordal pit.

this spinous pro− or absence of a facetted spinous process on the presacral ver− cess is absent (Evans and Klembara 2005.—Monjurosuchus sp. Four new char− . Monjurosuchus splendens and the Japanese additional material and associated specimens (frontal. an anteriorly elongated lachrymal. splendens. modification of the jugal. This requires some changes to the character Monjurosuchus differs from M. splendens (Gao et al. Evans and Klembara 2005). These differences probably distinguish the Japanese vation). In M. splendens in the length and codings for the genus in recent data matrices. 2000) to re− suchus. and the clavicle shape” (character number 63) was combined with position of the jugal/quadratojugal contact in relation to the another character “Clavicular facets on interclavicle” (char− postorbitofrontal/squamosal suture. Of the orbitofrontal and frontal facets. and A life reconstruction of the Japanese Monjurosuchus is postorbitofrontal (also closed in Hyphalosaurus. from the Lower facet. pre− choristodere share a unique combination of character states: maxilla and postcranial bones) would make the relationship closure of the lower temporal fenestra and corresponding clearer. Gao and Fox 1998). splendens at the species level but In summary. 15. RM personal obser− tebrae. tending over 1–2 tooth positions. such as those of width of the parietal and the relative proportions of the post− Evans and Klembara (2005) and Gao and Fox (2005). the Hecht 1992.340 ACTA PALAEONTOLOGICA POLONICA 52 (2). the level of ornamenta− acter number 65. quadratojugal. and Philydrosaurus). just be− tion on the posterior edge of the squamosal. Lazarus− shown in Fig. These similarities permit attribution of the Japanese cranial morphology and also extends the geographic range of choristodere to Monjurosuchus. and closure of the vertebral neurocentral sutures in the Cretaceous Tetori Group contributes new information on adult. but strongly tuberculate or spike−like projec− tions on the posterior margin of the squamosal (also Lazarus− Discussion suchus). and the presence low the postzygapophyses. a prefrontal con− tacting the maxilla anteriorly through an extremely short Phylogenetic position. 2007 Fig. 15. skull. using M. the length and ornamentation of the quadratojugal. process on the rear margin of the neural arch pedicel. numbers 1–73 were taken frontal and frontal facets in relation to the midline of the from Evans and Klembara (modified from Evans 1990. Life reconstruction of the Japanese Monjurosuchus. the level of the postorbito− characters used in the analysis. steeply sloping. However. The character of “Inter− presence or absence of a jugal/squamosal contact. squamosal. a short medial symphysis ex− construct the postcranial skeleton and soft tissue. the Japanese the genus. Monjurosuchus from M. a posteriorly constricted prefrontal that contributes to large medially expanded orbits.

and 64) and corrected the coding six missing entries in the original matrix (character numbers in three more characters (character numbers 14.pan. Cladograms showing alternative hypotheses of relationship for Choristodera based on a sequence of analyses described in the text. 67). by the authors) also added six missing data points (character For Monjurosuchus. Strict consensus of three most−parsimonious trees obtained after updating character codings (matrix of Gao and Fox 2005) for non−neochoristoderan taxa. New Monjurosuchus suchus. and 45) and required changes in two study thus changed codings in 17 characters for Monjuro− characters (character numbers 15.pl/acta52/app52−329. Strict consensus of four most−parsimonious trees obtained after adding codings for four new characters to the matrix used under C. B. 41. Cladogram obtained using the characters and matrix in Appendices 1 and 2. 53. 42. 38. 67. 25.MATSUMOTO ET AL. 63. This 11. Strict consen− sus of three most−parsimonious trees obtained after adding three new characters (from Gao and Fox 2005) to the matrix in Appendix 2. A. E.pdf . 72). 16. 60. The poorly known long−necked Chinese choristo− splendens specimens from the Yixian Formation (examined dere. C. acters (character numbers 74–77) were added (Appendix 1). has been included in the http://app.—CRETACEOUS CHORISTODERE FROM JAPAN 341 Fig. Cladogram obtained using the characters and matrix of Gao and Fox (2005) unchanged. Hyphalosaurus lingyuanensis. 56. D. the Japanese material provided data for numbers 1.

. The poorly known Pachystro− and 72) notionally support the relationship among Lazarus− pheus was excluded because it is too fragmentary and its sta− suchus. A 1993. tine. These placements agree with those of Similarly. with a one another and to Neochoristodera. upper temporal ours. Hecht 1992. Gao and Fox 1998). 73) are un− rescaled consistency index = 0. The first is that although Gao and Fox (2005) cite a count more than 16 (65). When we reran their analysis (using their choristodera. Finally. Lazarussuchus retains a basal position on the stem of (62). two characters were combined in our tree. Gao and Fox 1998. The more crownward position coded for Lazarussuchus. they did not run it in an analysis. palatal foramen preserved between pterygoid and pala− kawa) that Gao and Fox (2005) named Hyphalosauridae. In our analysis. Gao and Fox 1998. bootstrap support for the clade linking Philydrosaurus with the Jurassic Cteniogenys retains a basal position. the relative positions of Monjurosuchus and Lazarusuchus. consistency index = 0. and many and Hyphalosaurus. al. there were reduced pterygoid facet (25). Their trees. A maxi− (7. at most. (1999) and also on personal observation of new speci− deres other than Cteniogenys. and five common characters have addi− . Four characters (16. 16A).342 ACTA PALAEONTOLOGICA POLONICA 52 (2). cervical neurocentral sutures open trees. cervical neural spine much lower than dorsal neu− (Evans and Klembara 2005) that appeared as their manu− ral spine (75). Fig. articular level Choristodera. slender and horizontal. Monjurosuchus. although Gao and Fox (2005) included the prob− other recent studies (e. four or five firms the monophyly of Neochoristodera (Evans and Hecht characters firmly support a crownward placement of Laza− 1993) including Champsosaurus. However. two (29.g. and then Tchoiria were consis− matrix). prefrontal and frontal equal in length (72). 77) supporting the position of Lazarus− mum−parsimony analysis including all attributed choristo− suchus above Cteniogenys and Philydrosaurus.579. We ran a bootstrap analysis The relationships of the non−neochoristoderan taxa have on their consensus tree and found only 56% bootstrap sup− been more controversial (e. as the sister taxon of Monjurosuchus in a clade that is the sis− Consequently. Simoedo− russuchus in relation to Cteniogenys. However. of the 10 characters linking all a single shortest tree (length = 134. Ikechosaurus clade.g. cervical vertebra ences. A second difference is that we of Lazarussuchus in this cladogram contrasts with that of have coded more characters for Hyphalosaurus. while Philydrosaurus is the sister taxon of lower than mandibular symphysis level (74). 29. caudal neural spines long and nar− new description and phylogenetic analysis of Lazarussuchus row (70). amination of original material in the collections of the IVPP 2005). seven (58). and Tchoiria. without contribution from vomer (19). like contact (9). respectively) form a small clade charac− of the characters and codings are the same (see Appendix 2 terised by five synapomorphies: number of ossified carpal el− for details). All characters were equally Monjurosuchus and neochoristoderes. based on ex− previous studies where it is basal (e. a few characters differ in three additional trees in which the only difference was in (four new characters in the matrix of Gao and Fox [2005]. prefrontal−nasal contact) is difficult to code on The relationships of the Lower Cretaceous Asian choristo− crushed skulls. in total. mens by the senior author. Thus. Ksepka et lematic Pachystropheus in their data matrix (omitted by us). bootstrap analysis of our original tree shows only one deleted in theirs. 2005.0b10 (Swofford 2002) and and the last (72) shows state 2 in Lazarussuchus but state 1 in the exhaustive search mode. ptery− tions of Monjurosuchus and Hyphalosauridae in relation to goid process of quadrate low. The Japanese and Chinese long−necked taxa (Shokawa try (Evans 1990.g. 69. Of seven characters weighted and the tree rooted by outgroup (Youngina). dorsally directed orbits (10). 30. they did not incorporate the new data broad facets (76). the (35). Champsosaurus. However. only two (61. Running the analysis third difference is that while most of the characters are con− again but permitting one extra step had little effect. 30) are uncoded in Lazarussuchus. 36.. pterygoquadrate foramen present and enclosed by neo− positions of Lazarussuchus and Philydrosaurus. we obtained the same topology (Fig. Evans and Klembara 2005). resulting gruent between the two analyses. Ikechosaurus. Remov− tently placed as consecutive sister taxa of the Simoedosaurus– ing their additional outgroup (Araeoscelidia) had no effect. and neochoristoderes. saurus.. 2005. deran taxa (except the problematic Pachystropheus) yielded 11) are unequivocal. albeit sometimes with caveats (Evans and Hecht (resulting in a difference of 31/75 codings for this taxon). tooth enamel infolded at base two major differences between their phylogeny and ours. and four in ours). Ksepka et al. large. nasal/premaxilla to assess the position of their new genus. Ev− port for the clade excluding Lazarussuchus and only 59% ans and Klembara 2005. In their morph and quadrate (39). The problematic Lazarussuchus emerged in their matrix nor discuss the phylogenetic conclusions. 18/75 characters in their analysis are mis− ter group of Neochoristodera. but of these tus as a choristodere is unconfirmed.776. deres were analysed using PAUP 4. Chinese Philydrosaurus (Gao and Fox 2005) one node above Both matrices and character lists share a common ances− it. only two (7. with the Monjurosuchus. 2005). choristoderes except Cteniogenys. one (16. 2005). quadratojugal Their two most−parsimonious trees differed only in the posi− bears a cotyle meeting a rounded quadrate process (24). cervical postzygapophyses horizontal with script was in revision. found a monophyletic Neochoristodera and a small fenestra posteriorly flared and substantially larger than orbit clade of long−necked choristoderes (Hyphalosaurus + Sho− (13). Neochoristoderans share 12 synapo− Gao and Fox (2005) also ran an analysis of Choristodera morphies: elongated and fused nasals (8). This tree con− equivocal for Lazarussuchus. Within Neo− Monjurosuchus. there are a number of important differ− ements reduced to. Gao and Fox 1998. 16B). 2007 phylogenetic analysis based partly on the description by Gao weak support (58%) for the clade encompassing all choristo− et al. 11–13.

sug− Evans and Klembara 2005).pl/acta52/app52−329. for Monjurosuchus. for Laza− Evans and Klembara 2005). Gao and Fox 1998. In order to determine the effect of the coding differences Gao and Fox (1998. On the other hand. Phylogenetic tree for Choristodera plotted against time. Finally there were several in− might be a derived taxon with some size−linked character re− stances. We lation to other choristoderes remains unresolved.—CRETACEOUS CHORISTODERE FROM JAPAN 343 Fig.MATSUMOTO ET AL. the relationship between Lazarussu− Lazarussuchus is problematic.pan. 17. sence of the parietal foramen (3). although Evans and Hecht (1993) gests that Lazarussuchus shares at least three synapomor− also offered an alternative hypothesis that Lazarussuchus phies with all other choristoderes: absence of the parietal fo− http://app. where there is disagreement on the coding of character one extra step was enough to yield a subset of trees that moved states (see Appendix 3). 16E). Laza− chus and other choristoderes was supported by two charac− russuchus has been placed at the base of the choristoderan tree ters. and ab− (Hecht 1992. for all non−neochoristoderes (Fig. In The phylogenetic position of the Oligocene–Miocene Gao and Fox (1998). russuchus. Similarly. particularly for those taxa outside of Neochoristo− versals. such is the also reran our analysis after adding their additional charac− ambiguity of its position (moving up or down the tree as ters (Fig. for all The overriding conclusion from the set of analyses pre− non−neochoristoderes and Tchoiria. prefrontal elongated with median contact (2). 16D). Evans and Klembara (2005) showed that dera. 2005) placed Lazarussuchus as the sister on the resulting trees. and after adding our sented here is that while the position of Lazarussuchus in re− four new characters to their recoded matrix (Fig. excluded from Choristodera (contra Gao and Fox 2005). Lazarussuchus from the basal position. This study. 16C). The results of these analyses are described in characters are added and codings changed) that it cannot be more detail in Appendix 3. In all recent studies. we performed a series of separate anal− taxon of Choristodera. however. tional or slightly altered states.pdf . because Lazarussuchus seemed to have yses on the Gao and Fox (2005) matrix: after updating the none of the derived characters diagnosing the group (but see codings for Hyphalosaurus. Evans and Manabe 1999.

Ja− matic character. and Shizuo Shimojima (Shokawa Town. then closure of the lower temporal fenestrae is an enig− ture. chus. and Yuan Wang (IVPP) for suchus. parietal. This is known saurus.67) than does based mainly on differences in the morphology of the verte− Lazarussuchus (0. The third type is characterized by a short snout. and 63) support a placement of Lazarussuchus closed in the ancestry of post−Cteniogenys choristoderans above Cteniogenys. XingHe Liu (DR). and new specimens of Monjurosuchus. However. In terms of 57. Lazarussu− (17). atal and braincase characters. Zhonghe Zhou. dorsal flange of maxilla low. Hakusan City. and an upper Hirano (Waseda University. Meeman Chang. with the caveat that the nothing is known of the Formation (Upper Hauterivian–Barremian) of northeastern skull of the Japanese Shokawa. 12.344 ACTA PALAEONTOLOGICA POLONICA 52 (2). The second skull ing it available for study. with dorsal border have been fully diapsid. and probably represent a new species. The new Monjurosuchus from the Kuwajima and based solely on postcranial characters. . Japan. Tokyo. and the latter taxon tends to plot in a more basal position. 3: interorbital contriction of the frontals) is problematic since it is inti− Choristoderan bones from the Lower Cretaceous Kuwajima mately related to eye size. Gifu Prefecture. opened in Cteniogenys and neochoristoderes. Philydrosaurus has a skull in− History. or it ters 8. A second pur− Conclusions ported synapomorphy (Gao and Fox 2005.56. Gunma. the basal position of the Oligocene–Mio− zoic representative of the Lazarussuchus lineage would be cene Lazarussuchus is not confirmed. 2007 ramen (3). and vomer−maxilla contact present pendently in Philydrosaurus. nor is a relationship illuminating. 11. This needs to Support for a monjurosuchid clade incorporating both be tested with a broader analysis of diapsid relationships. squamosal. 52. the latter hypothesis is more lating to fusion of the neurocentral sutures. and Monjuro− Fig. and reopened early in neochoristoderan history. Gifu Prefecture. particularly with respect to pal− and also our knowledge of its morphological variation. Monjurosuchus was known only from the Yixian both studies. Ishikawa Prefecture. 51. although characters re− the number of required changes. large dorsally directed orbits. We thank Tatsuya Sakumoto and Yoshinori type is characterized by closed lower temporal fenestrae.—The choristoderan skull shows many unique features amongst Reptilia and is particularly characterized by dorsoventral compression and posteriorly expanded supra− temporal fenestrae. The mayors and administra− closed temporal fenestrae of the other taxa. Japan) for helpful discussion. Japan) for kindly arranging access to new material of Monjurosuchus and Hyphalosaurus during a temporary exhibition. fusion of the sa− parsimonious but it will be tested by subsequent finds. Implications. 15. and such is the instability of Our cladistic analysis supports the monophyly of Neo− the lower part of the tree that new data may well support al− choristodera and of the Hyphalosauridae of Gao and Fox ternative hypotheses of relationship. If the tree is cor− tions of Kuwajima District (Shiramine. the termediate between the second and third types in that the Wondrous World of Dinosaurs. Monjurosuchus. temporal fenestra that is equal in size to the orbit. 0. The monophyly of Hypha− brae. When the matrix of Gao and Fox (2005) is updated. coded at 0). Simoedosaurus. Japan). Recovery of a Meso− (2005). orbital width divided by the posterior width. Japan) and Shokawa Town (Takayama City. only in the basal Cteniogenys and may represent the ancestral Yoshikazu Hasegawa. However. Takayama City. between Philydrosaurus and Monjurosuchus. five unequivocal characters (2. and two skull shape is similar to that of Cteniogenys but it shares the anonymous referees for their comments. Japan. Only one charac− ter (Gao and Fox 2005. have been generous in their hospitality and support. even within Choristodera. Acknowledgements skull morphology is diverse and can be classified into three types. and Tchoiria). Ishikawa Prefec− rect. and arranging access to holotype specimens of Ikechosaurus and Hyphalo− by elongated and more laterally facing orbits. Hiromichi shorter snout. Japan) for drawing the reconstruction in shown by Hyphalosaurus. Lazarussuchus. Previ− losauridae and of Neochoristodera is strongly supported by ously. may be attributed to the Jehol (coded as 1). Ishikawa Prefecture and the Okurodani Forma− the width of the posterior part of the frontal. A cral and caudal ribs to their vertebrae. Okurodani formations (Berriasian–Hauterivian) of Japan ex− It is very obvious that additional data are needed for all tends both the geographic and temporal range of the group non−neochoristoderan taxa. Philydrosaurus tion. Monjurosuchus and Philydrosaurus is weak. so that the relationship is China. and greater develop− more radical alternative would be that the lower temporal ment of the flexor tubercle of the unguals could be linked to fenestra was closed in the ancestor of choristoderes and more terrestrial habits. at Chiba Prefecture. and jugal. 75: the presence of a distinct process on the ischium) supports the relationship. the lower fenestra either closed inde− inflected medially (5). and the ancestor of a clade containing Hyphalosaurus four independent and unequivocal characters (their charac− and Shokawa (assuming the latter shows this feature). 61) support the reversed position. and an open lower temporal fenestra. actually has a broader interorbital frontal (inter− genus Monjurosuchus. Ikecho− tion. quadratojugal. a Kobayashi (SBEI). small dorsally The material described here was collected by Ichio Yamaguchi and pre− directed orbits. and Yuji Takakuwa (Gunma Museum of Natural morphology for Choristodera. itself a character and depends on Formation. and we are grateful to them for mak− saurus. One is characterized by a long−snout. Japan) for arranging access to specimens and loans. Assuming the choristoderan ancestor to pan). This pared by Mikiko Yamaguchi (formerly of Shiramine Board of Educa− type is found in Neochoristodera (Champsosaurus. This type is Utako Kikutani (Kanagawa.

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is added. (2000) code Monjurosuchus (0) for this char− strong posterolateral wing−like expansion (2). Lachrymal perforation: perforated by one or two foramina 27. subtriangular tracted close to the midpoint of the marginal tooth row (1). anterior to the condyle (1). but in the well−preserved M. splendens (DR0003C) and 33. pro− 1. anterior to suborbital fenestra (0). opening at the junc− ans and Klembara (2005) and Ksepka et al. more than 25. Vomer−maxilla contact: absent (0). Preorbital skull proportions: moderately short and rounded meeting a rounded quadrate process (1). re− 37. Pterygoid flange: consisting of pterygoid only and ventrally acter codings for Monjurosuchus have been changed in sev− directed (0). Shape of suborbital fenestra: narrow. moderate postero− length to orbit (1).pdf . 14. Dorsal process of jugal: as prominent as anteroventral process riorly from the opening. Evans anteriorly and posteriorly by pterygoids (2). elongate but less than 50% of skull length (1). Basal infolding of tooth enamel: absent (0). low. (1). the occipital condyle (0). lapping facet for the quadrate (0). 15. 6. with a 5. long midline suture (1). present (1). palatine. nearly terminal (0). and Evans and Manabe (1999). 10. between pterygoid and fied from Evans (1990). Prefrontals: anterolateral and separated by frontals (0). a large pterygoid facet (0).pan. 32. present (1). close to the premaxilla (0). Hecht (1992). External nares: paired. but character 64 has been 20. paired.. elongate and paired (1). Basal tubera of braincase: weakly developed (0). large and dorsally directed (2). Lachrymal: enters external narial opening (0). orbital margin (0). 22. Internarial: absent (0). slender. Enclosure of interpterygoid vacuity: enclosed anteriorly by orbital margin (1). prominent but much shorter than anteroventral process (1). elon− cess and cotyle are reduced and the two bones are sutured (1). Marginal tooth sockets: circular (0). Monjurosuchus sp. nasals intervene between premaxillae 29. with dor− reduced pterygoid facet (1). present (1). duced as a small triangular bone (2). small and near posterior bor− 11. but open posteriorly (0). and Klembara (2005) code Monjurosuchus (0) for this character. dis− with straight medial edge (1). short. nasals do not contact premaxillae (2). Char− 21. vertically oriented with 50% of skull length (2). median contact (1). on snout (0). Parasphenoid/pterygoid contact: no contact (0). Gao et al. (SBEI 1792) and M. Pterygoid process of quadrate: broad. Palatal teeth: palate covered by shagreen of small teeth (0). oval.MATSUMOTO ET AL. between premaxillae (0). acter. elongate (0). gate and dorsally placed (1). Upper temporal fenestra: shorter than orbit (0). 24.pl/acta52/app52−329. Squamosal/parietal suture in occiput: near posterior end of 35. low. Basipterygoid/pterygoid joint: basipterygoid process articu− lates with pterygoid cotyle as part of a metakinetic joint (0). terminal and confluent (2). retracted poste− 26. enclosed anteriorly by 12. lachrymal foramen between lachrymal and prefrontal (1) 28. splendens (DR0003C) 16. 18. 38. roughly equal in (0). modified and four new characters (74–77) are added. enclosed both elongate to middle level of inferior temporal opening (1). kidney−shaped (2). Nasopalatal trough: absent (0). midline anteriorly (0). Parietal: having broad contact with the postorbital/postfrontal 34. posteriorly flared and substantially longer than lateral expansion (1). but remains elongate (1). through entire length of element (2). One new taxon. Palatal foramen: absent with vomer. palatine and pterygoid as part of external wall of braincase and medial wall of temporal http://app. 36. Gao and Fox (1998). tight contact or fusion (2) contact long. placed far back (2). elongate paired pterygoid tooth batteries separated by nasopalatal trough and fused (2). broad V−shaped contact at dorsal midline (1). the two bones are fused (2). long. Orbits: large and laterally directed (0). median 23. Neomorph (sensu Fox 1968) in braincase: absent (0). Size and location of interpterygoid vacuity: large and extends rected (1). postfrontal and postorbital fused (2). small and dorsally di− 30. Shape of parasphenoid: isosceles triangle with long rostrum 13. 7. Location of choana: anterior. quadratojugal bears a cotyle 4. Location of craniomandibular joint: roughly on a level with narrow wedge (2). the condyle (2). present 19. Nasals: short and paired (0). Nasal/prefrontal contact: straight line contact dorsolaterally (0). Dorsal flange of maxilla: high and vertical (0). curs in anterior part of tooth row (1). Midline contact of pterygoids: separate or just touching in the (1). consisting of pterygoid and ectopterygoid and hav− eral places to reflect new observations and interpretations of ing a horizontal overlap (1). Quadratojugal/quadrate articulation: having a simple over− 3. Nasal/premaxilla contact: nasals contact but do not intervene (2). 2. (0). transverse expansion oc− complex (0) having reduced contact (1). broader anteriorly with short rostrum. Hyphalosaurus. Monjurosuchus sp. Dorsal process of squamosal: broad and short (0). narrow rows of palatal teeth separated by nasopalatal trough 9. and horizontal. the upper temporal fenestra is roughly equal expanded laterally (1). half way along fenestra (1). moderately Monjurosuchus sp. wing−like (2). slender anteriorly with long rostrum and orbit (2). Postfrontal/postorbital fusion: bones discrete and both enter der of the fenestra (1). bones discrete but postorbital excluded from 31. without contribution from vomer (2). little or no process (2). Parietal foramen: present (0). (0).—CRETACEOUS CHORISTODERE FROM JAPAN 347 Appendix 1 The characters used in the analysis are taken mainly from Ev− meeting at a closed three−point suture (0). absent in superior temporal fenestra (0). (2005) and modi− tion between the three palatal bones (1). 8. further re− (0). the process is oriented vertically in sal border inflected medially (1). strongly expanded posterolaterally and in size to the orbit (1). posterior to 17. slender and pterygoid and posteriorly by parasphenoid (1). the authors. absent (1). pterygoids. clear sutural contact limited to posterior two−thirds of length (1). (0). present (1).

almost as 48. platyan with notochordal canal closed (1). Hyphalosaurus have an exceptionally low neural spine. 52. physes: absent (0). Prefrontal−frontal lengths: prefrontal shorter than frontal (0). Bone structure: cancellous (0). 49. moderate elon− gle to the horizontal in adults (0). Paroccipital process/quadrate contact: only tip of process reduced (1). Lower temporal fenestra: open. the symphysis is small and terminal in Monjurosuchus sp. (0). shorter 45. T−shaped and clavicle facets continue across the 42. 66. caudal ribs fused (1). Monjurosuchus sp. in width (0). distal end flared (1). but M. reduced to. in Monjurosuchus. supra− ing distally (1). longer (1). in Lazarussuchus. Iliac blade. 55. 76. low or no keels boundaries (0). short and jurosuchus (DR0003C) and Lazarussuchus. amphi− 72. 77. half the length of the tooth row with longer inclusion of splenial 63. inter− occipital lightly arched. taper− 44. (SBEI level (0). (0). Caudal neural spines: low (0). robust. Supraoccipital/parietal contact: free from parietal (0). cylindri− 46. these cervical zygapophyses have expanded 56. ungual: distinct and well developed (0). elongated process lies in a ity (1). flexor tubercle low and 40. Number of ossified carpal elements: at least nine (0). Presacral vertebral centra: longer than wide (0). they are vertical in Monjurosuchus splendens. Number of sacral vertebrae: two sacrals (0). the midline (0). flask−shaped. pachyostotic in adult (2). strongly elongate. 69. 67. in Shokawa and (2). Dorsal neurocentral sutures: closed in adult (0). parasphenoid and opens ventrally (1). 2007 fossa (1). splendens. Sacral and caudal ribs: fused to the vertebra in the adult (0). 1792) has small processes. orbitofrontal. open (1). Interclavicle shape and clavicular facets on interclavicle: (2). Lazarussuchus and Monjurosuchus (1). facets (1). Fibula shape: proximal and distal ends of similar width. Articular and mandibular symphysis level: roughly equal present and bear accessory facets (2). Gastralia thickness: thin. Monjurosuchus has cylindrical sphenoid (1). articular level is lower than mandibular symphysis (1). post− (1). (SBEI 1792) has a facet for the 58. the than high (1). lightly built (0). Naso−maxillary contact: nasals contact maxillae laterally (0). absent through loss (1). longer than frontal (1). Flexor tubercle or process on the ventral surface of the by neomorph and quadrate (1). derived condition in Hyp− 53. Orientation of paroccipital processes: horizontal (0). 75. free odontoid process unfused to axis (1). groove is flanked by deep ventral (1). splendens (DR0003C) derived condition in Neochoristodera. character 63 (in Evans and Klembara 2005). long and narrow (1). Cervical neurocentral sutures: closed (0). 1223. the prefrontals are spool−like (1). stem shorter than lateral processes (2). centra (1).. Vertebral centrum: amphicoelous and notochordal (0). splendens (IVPP V13279). strongly deflected ventrally (2). prominence of axis (0). Atlas−axis complex: little or no development of anterior odontoid thick as axial ribs and pachyostotic (1). much lower than dorsal neural spine (1). Cervical vertebral centra length: longer than high (0). or proxi− trate parasphenoid and opens ventrolaterally (0). 61. little inclusion of splenial (1). Dorsal vertebral centrum shape: subcylindrical (0). . Cervical neural spine: equal in height to dorsal neural spine 54. in M. supraoccipital keeled. Caudal zygapophyses: lie at a small angle to the horizontal (0). Posterior opening of internal carotid artery: does not pene− 68. the femur has trough of the quadrate that is lined by a thin sheet of the been sectioned and the bone structure is cancellous (0). and parietal facets extend 65. and quadratojugal (1). the flanges (1). elongated process lies in a trough formed by 60. parietal facet is wider than prefrontal facet caudal blood vessels (0). equal in length to frontal (2). does not have the process (0/1). blade expanded. dorsal margin: essentially vertical or at a steep an− 41. more than 16 (1). Posttemporal fenestra: present (0). present and enclosed 59. remain open 71. 50. nasals reduced and separated from maxillae by prefrontals (1). Small spinous processes below the presacral postzygapo− halosaurus. Cervical vertebral count: eight or nine (0). Pila antotica: remains unossified (0). slender and essentially parallel−sided depressed (1). (1). proximal end narrow. Frontal: prefrontal and parietal facets are equal. nearly vertical (1). 39. Pterygoquadrate foramen: absent (0). Centra of caudal vertebrae: bears shallow ventral groove for equal. meets quadrate (0). 47. parietal facet is wider than the prefrontal facet. three sacrals (1). anteriorly (1). interclavicle shape is rhomboid and clavicles meet at an angle in (SBEI 1792) and M. equalling at least 62. Interclavicle stem: long. having reduced medullary cav− the quadrate and neomorph (1). the interclavicle is T−shaped of mandible (0). Char− (1). number neomorph on the quadrate (1). Monjurosuchus sp. often triangular (1). squamosal. Mandibular symphysis: small and terminal (0). present but without accessory facets (1). dorsal margin essentially hori− gation equalling length of one third or less of the tooth row with zontal. Shokawa and four sacrals (2). broad in anterior and central portions. 74. slightly 64. horizontal with broad facets (1). Cervical zygapophyses: project dorsolaterally with narrow sacral ribs free. Hyphalosaurus and Monjurosuchus. at most. closed by apposition of jugal. 43. exposed on ventrolateral surface of mandible and the clavicular facets continue across the midline (1). ossified as part of the cal (circular cross−section) (1). in Mon− 51. with complete or incomplete brae are strongly keeled like the cervicals (0). 70. splenial confined to medial side in Monjurosuchus sp. to posterior margin (2). sacral and caudal ribs free facets (0). Hyphalosaurus. fenestra is absent in Monjurosuchus sp. or nearly 57. penetrates mally wider (0). in adult (1). neomorph (2). Ventromedial crest of dorsal vertebra: anterior dorsal verte− 73. seven (1). with sutural surfaces for parietal placed clavicle stem is long and slender in Monjurosuchus (0). acter 65 (in Evans and Klembara 2005) was combined with SBEI 1792 (0).348 ACTA PALAEONTOLOGICA POLONICA 52 (2). Lateral exposure of splenial: splenial confined to medial side midline (1).

—CRETACEOUS CHORISTODERE FROM JAPAN 349 Appendix 2 A = 0/1.pdf .pan. B = 1/2. C = 0/2 5 10 15 20 25 30 35 40 Youngina 00000 00000 00000 00000 00000 00000 00000 00000 Lazarussuchus 11111 21002 2?100 211?0 ???0? 1???? ???00 ?0??? Cteniogenys 22111 10A20 0?A00 ?1110 11100 10000 10000 ?01?? Shokawa ????1 ????? ????? ????? ????? ????? ???0? ????? Champsosaurus 22121 21211 B1200 21221 12211 11211 22201 12112 Tchoiria 221B1 112?1 B1210 21120 11111 11111 11111 2111? Ikechosaurus 22111 1?211 21211 1?120 11111 21111 11111 2111? Simoedosaurus 22111 11211 21211 11120 11111 21111 11111 2111? Hyphalosaurus 02101 1?002 2?10? 011B0 ????? B??00 ???00 1???? Monjurosuchus 02101 1?002 220A0 1???? ??B00 11111 1??00 1?100 Philydrosaurus 1?111 10002 00000 0???? ????? 1???? ???00 01??? 45 50 55 60 65 70 75 77 Youngina 00000 00000 00000 00000 00000 00000 00000 00 Lazarussuchus 11??? ???00 00A20 11000 101?0 00100 12100 01 Cteniogenys 000?? 00111 00012 11?11 00??? 00?00 0000? ?0 Shokawa 00??? ???11 0?011 1111B 00001 01111 ????1 1? Champsosaurus 21210 00111 01012 11?11 11110 A1110 01010 00 Tchoiria 20111 ?1111 0111? ?1??B 11110 011?0 ?101? ?0 Ikechosaurus 10121 11?11 11112 ?0?1? 11100 0A110 ?1010 00 Simoedosaurus 10121 11111 11112 00??2 11110 11110 01010 00 Hyphalosaurus ????? ????1 01?12 ?110B 10011 01111 00101 11 Monjurosuchus 00??? ???01 0?A10 10010 10100 01100 02100 01 Philydrosaurus 0???? ??101 01011 11??? 10100 ??100 0010? ?0 http://app.MATSUMOTO ET AL.pl/acta52/app52−329.

19 [17] ? [1].684) in which Lazarussuchus. 34 (4) We then updated codings for all non−neochoristoderan taxa in the [32] 0 [?]. 60 [58] ? [1]. 27 [25] ? [0].350 ACTA PALAEONTOLOGICA POLONICA 52 (2). 47 [45] 1 [?]. 43 [41] 1 [0]. RC = 0. Hyphalosauridae as the sister group of Neochoristodera. 47 [45] 1 [?]. 58 [56] ? [1]. The reanalysis resulted in Analyses three shortest trees (L = 142. each with a very differ− [0]. 61 [59] ? [1]. 7 [6] ? [1]. We obtained four shortest Tchoiria: 33 [31] 1 [1/2]. 20 [18] ? [1]. 60 [58] 1 [?] were the most basal choristoderes. 21 [19] ? [1/2]. RC = 0. 62 [60] ? [1/2]. 60 [58] 0 [1] rus. Differences in coding of character states and Monjurosuchus and Philydrosaurus forming a clade). 57* [55] 1 [2] (Monjurosuchus + Philydrosaurus) is supported (75%) and lies (4/75) at the base of Choristodera. 26 [24] ? [0]. 12 [11] ? [2]. clade comprising all remaining taxa (75%). Key: in each line. 54 [52] ? [1]. 22 [20] ? [0]. 63 [61] 0 [1]. 9 [8] 1 [0]. 20 larger.605). the strict consensus tree Other choristoderes (Fig. 61 [59] ? [1]. 22 [20] 0 [?]. 30 [28] 0 [?]. represents the code in our suchus formed consecutive sister taxa to Neochoristodera (i. 11 [10] 0 [1]. 31 [29] 0 [?]. 63 [61] 0 our four new characters. 64 [63] ? [0]. 29 [27] ? [1]. in which Lazarussuchus and Hyphalosaurus: 1 [1] ? [0]. 69 [67] ? [1] (32/75) Philydrosaurus: 8 [7] ? [0] (state description differs in original tree. that is. the fourth. 12 [11] 0 [2]. Updating the codings for Tchoiria ? [0]. 33 [31] ? [1]. 60 [58] 1 [?] 62 [60] 2 [1] (17/300) relation to Neochoristodera. 13 [12] 2 [0]. 18 [16] 0 [1]. Five charac− (length [L] = 126. 59 [57] ? [1]. 34 [32] 1 [1/2]. 16 [14] ? [0]. 50 [48] ? [1]. suchus. 65 [64] ? [0] (12/75) 147. (put as ?). 15 [73] ? [1]. two of Monjurosuchus: 8 [7] 0 [?]. 31 [29] tree (Fig. 68 [66] ? [0]. and the Hyphalosauridae changed position in [57] 0 [1]. 42 [40] ? [0]. Champsosaurus: 12 [11] 1 [1/2]. 16C) supports only Neochoristodera and Hyphalosauridae Shokawa: 51 [49] ? [1]. and finally Monjuro− trix. above them Cteniogenys. 57* [55] 1 [2]. 51 [49] 0 [?]. CI = 0. 9 [8] ? [0]. 19 [17] ? supported a relationship between Monjurosuchus and Philydro− [1]. RC = 0. 66 are exactly the same. 43 [41] 0 [?].69). 65 [64] ? [0]. An asterisk marks where an additional state has been added or the relationship between Philydrosaurus and Monjurosaurus a state changed. Lazarussuchus in a basal Four characters are new in each matrix. 69 [67] ? [1] (31/75) Gao and Fox’s (2005) new characters. 59 [57] ? [0]. 53 [51] ? [0]. position.e. CI = 0. Monjurosuchus. CI = 0.757. 13 [12] 2 [0]. the second number in the square bracket and Fox's (2005) matrix and obtained a single shortest tree (L = is the equivalent character number in Appendix 1. 32 [30] ? [0]. 17 [15] ? [0]. 53 [51] 1 [0]. 55 [53] 0 [0/1]. except in breaking order to understand the effect of modifying the character state the weak relationship between Monjurosuchus and Lazarus− codings between the two matrices. matrix. 32 [30] ? [1]. 64* [63] ? [1] (18/75) in three shortest trees (L = 136. Lazarussuchus: 8 [7] 0 [1]. their charac− Total difference (101/450) ter 3. forming the sister group to a second. 2007 Appendix 3 Basic data (1) We updated only the codings for Hyphalosaurus (31/75) in Gao The matrix in Appendix 2 has 77 characters. 22 [20] 0 (5) In the final analysis on Gao and Fox's (2005) matrix. only one of the four [0]. 17 [15] ? [0]. 18 [16] ? [0]. (6) The last analysis was a rerun of our matrix incorporating three of 65 [64] ? [1]. Again. the matrix of Gao and and Fox's (2005) matrix. 25 [23] ? placed Lazarussuchus above Cteniogenys. The strict consensus tree (Fig.. rescaled consis− ters are almost the same with slight changes in character states. 63* [61] 0 [1].541) with rela− The following maximum−parsimony analyses were performed in tively little difference from our original tree. 38 [36] 0 [?].589). 44 [42] 0 [1]. 41 [39] the base of Choristodera.812. 57 (based mainly on Ksepka et al. in a polytomy with remaining taxa. 2 [2] ? [2]. 53 Cteniogenys. 70 as clades. we added in [?]. 38 [36] 0 [1]. The result was a single shortest tree Fox (2005) has 75. 10 [9] ? Cteniogenys were each basal in two. 57* [55] 1 [2]. again in square brackets. 28 [26] ? [1/2]. 58 [56] 1 [0] [?]. 53 [51] ? [0]. CI = 0. text). 68 [66] ? [0]. 16E) has a polytomy of [29] ? [0].754. Philydrosaurus. ei− Ikechosaurus: 43 [41] 2 [1]. 59 Lazarussuchus. Hyphalosauridae. 2005) had little effect. 31 saurus. The strict consensus [1/2]. The fourth. RC = 0. The final figure in bold shows the total number of was not supported). 40 [38] ? [1]. 53 [51] 1 trees (L = 127. the third number 128. was omitted as it is problematic. 63 [61] 0 [1]. character codings that differ for that taxon or group of taxa. In all cases. 35 [33] 0 [?]. (3) Updating only the codings for Lazarussuchus in Gao and Fox's Neochoristodera matrix (18/75) had a greater effect. Lazarussuchus. 51 [49] ? [1]. matrix of Gao and Fox (2005) and reran the analysis. . 38 [36] 0 [1]. 57* [55] 1 [2]. 28 [26] ? [1]. However. CI = 0. 9 [8] 1 [0] (proportional).819.825. 30 [28] ? [1]. In a 50% majority [68] 0 [1] (5/75) rule consensus tree. [18] 0 [1]. Two tency index [RC] = 0. This resulted in four shortest trees (L = [1]. 26 [24] ? [0]. and the third 16 [14] 0 [0/1]. 9 [8] ? [0]. with Hyphalosauridae and Neochoristodera in the crown.707) with a topology essentially unchanged further characters are the same but were combined in our matrix. 59 [57] 1 [0] ent topology from that in Gao and Fox (2005). 61 [59] ? [0]. consistency index [CI] = 0.748. 16D) has Lazarussuchus and Cteniogenys unresolved at ? [1]. Gao and Fox's (2005) Monjurosuchidae Cteniogenys: 19 [17] 0 [1]. and Philydrosau− [51] ? [0]. 38 [36] 2 [0/2]. 55 [53] 0 [0/1]. 64 weaken bootstrap support in the non−neochoristoderan part of the [62] ? [1]. 57* [55] 1 [2] ther Monjurosuchus alone or Monjurosuchus + Philydrosaurus Simoedosaurus: 42 [40] 1 [?]. RC = 0. except to [55] 1 [0]. Of these. 62 [60] ? [0]. the first number is the character number in the ma− (2) We updated only the codings of Monjurosuchus (32/75) in Gao trix of Gao and Fox (2005). from that of Gao and Fox (2005). 44 [42] 0 [?]. which had Lazarussuchus in the basal position. This resulted 59 [57] 0 [1]. Ctenio− or symbol represents the code for that character in Gao and Fox's ma− genys.