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Appropriate large-scale citizen-science data present important new opportunities for biodiversity modelling, due in part to the wide spatial coverage of information. Recently proposed occupancy modelling approaches naturally incorporate random effects in order to account for annual variation in the composition of sites surveyed. In turn this leads to Bayesian analysis and model fitting, which are typically extremely time consuming. Motivated by presence-only records of occurrence from the UK Butterflies for the New Millennium data base, we present an alternative approach, in which site variation is described in a standard way through logistic regression on relevant environmental covariates. This allows efficient occupancy model-fitting using classical inference, which is easily achieved using standard computers. This is especially important when models need to be fitted each year, typically for many different species, as with British butterflies for example. Using both real and simulated data we demonstrate that the two approaches, with and without random effects, can result in similar conclusions regarding trends. There are many advantages to classical model-fitting, including the ability to compare a range of alternative models, identify appropriate covariates and assess model fit, using standard tools of maximum likelihood. In addition, modelling in terms of covariates provides opportunities for understanding the ecological processes that are in operation. We show that there is even greater potential; the classical approach allows us to construct regional indices simply, which indicate how changes in occupancy typically vary over a species’ range. In addition we are also able to construct dynamic occupancy maps, which provide a novel, modern tool for examining temporal changes in species distribution. These new developments may be applied to a wide range of taxa, and are valuable at a time of climate change. They also have the potential to motivate citizen scientists.

Beishanodon, new genus TYPE SPECIES: Beishanodon youngi, new species. ETYMOLOGY: After the Beishan Hills, in which the type locality occurs. DIAGNOSIS: As for the type and only known species.

Beishanodon youngi, new species HOLOTYPE: PKUP V3007, incomplete skull with well-preserved snout and left temporal region, but lacking the right zygomatic arch and much of the occiput; only known specimen. ETYMOLOGY: The specific epithet is in honor of the late Chinese paleontologist C.C. Young, who named and described the first cynodont known from China. TYPE LOCALITY AND HORIZON: Beishan locality, Quarry-3, in Beishan Hills (41 ° 56 9 739 0 N/ 96 ° 32 9 023 0 E), northern Gansu Province, China; Lower Triassic dark shales of the Hongyanjing Formation (see above discussion). DIAGNOSIS: An Early Triassic sinognathine trirachodontid differing from the closely related Sinognathus gracilis by the following derived characters: triangular lateral process of frontal closely approaching but not entering dorsal rim of orbit; jugal/postorbital suture located at midlength of postorbital bar; short intertemporal crest, about one-half length of enlarged temporal fenestra; posterior border of osseous secondary palate straight; incisor roots buccolingually expanded; upper canine obliquely positioned lateral to paracanine fossa; upper postcanine teeth eight in number, and less expanded laterally; transverse crest in upper postcanines located posteriorly; maxillary tooth row terminating at level close to anterior border of orbit; smooth curvature from snout to zygomatic arch; major palatine foramen penetrating palatine at midlength of the element.

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)., Published as part of Gao, Ke-Qin & Fox, Richard C., 2005, A new choristodere (Reptilia: Diapsida) from the Lower Cretaceous of western Liaoning Province, China, and phylogenetic relationships of Monjurosuchidae, pp. 427-444 in Zoological Journal of the Linnean Society 145 (3) on page 439, DOI: 10.1111/j.1096-3642.2005.00191.x, http://zenodo.org/record/4687363, {"references":["Cope ED. 1876. [1877]. On some extinct reptiles and Batrachia from the Judith River and Fox Hills beds of Montana. Proceedings of the Academy Natural Sciences of Philadelphia 28: 340 - 359.","Evans SE. 1989. New material of Cteniogenys (Reptilia: Diapsida) and a reassessment of the systematic position of the genus. Neues Jahrbuch fur Geologie und Palaontologie, Monatshefte 1989: 577 - 589.","Evans SE. 1990. The skull of Cteniogenys, a choristodere (Reptilia: Archosauromorpha) from the Middle Jurassic of Oxforshire. Zoological Journal of the Linnean Society 99: 205 - 237.","Gao K, Evans S, Ji Q, Norell M, Ji S. 2000. Exceptional fossil material of a semi-aquatic reptile from China: the resolution of an enigma. Journal of Vertebrate Paleontology 20: 417 - 421.","Gao KQ, Tang Z, Wang X. 1999. A long-necked diapsid reptile from the Upper Jurassic / Lower Cretaceous of Liaoning Province, northeastern China. Vertebrata Palasiatica 37: 1 - 8.","Evans SE, Manabe M. 1999. A choristoderan reptile from the Lower Cretaceous of Japan. Special Papers in Palaeontology 60: 101 - 119.","Gao KQ, Fox RC. 1998. New choristoderes (Reptilia: Diapsida) from the Upper Cretaceous and Palaeocene, Alberta and Saskatchewan, Canada, and phylogenetic relationships of Choristodera. Zoological Journal of the Linnean Society 124: 303 - 353.","Ksepka DT, Gao K-Q, Norell MA. 2005. A new choristodere from the Cretaceous of Mongolia. American Museum Novitates 3468: 1 - 22.","Evans SE, Klembara J. 2005. A choristoderan reptile (Reptilia: Diapsida) from the lower Miocene of northwest Bohemia. Journal of Vertebrate Palaeontology 25: 171 - 184.","Hecht MK. 1992. A new choristodere (Reptilia, Diapsida) from the Oligocene of France: an example of the Lazarus effect. Geobios 25: 115 - 131."]}

PHILYDOSAURUS PROSEILUS SP. NOV. (FIGS 2���5) Etymology: Philydros + sauros (Gr.), water-loving lizard, referring to the presumed aquatic life style of the animal; proseilos (Gr.), meaning toward the sun, similar in meaning to the place name Chaoyang. Holotype: PKUP V2001, nearly complete skull and lower jaws in association with partial postcranial skeleton. Type locality and horizon: China, western Liaoning Province, Shangheshou near the city of Chaoyang; Early Cretaceous Chiufotang Formation. Known distribution: China, western Liaoning Province, Early Cretaceous Chiufotang Formation. Diagnosis: A medium-sized monjurosuchid choristodere sharing with Monjurosuchus the following derived character states: frontal markedly narrow and hourglass-shaped; anterior extension of lacrimal close to nasal/prefrontal suture; supratemporal fenestra small; infratemporal fenestra closed by expansion of surrounding elements, including postorbital, quadratojugal and squamosal. Differing from Monjurosuchus in having a more elongate skull; elongate narial openings and orbits; presence of a ���supratemporal trough��� formed by the postorbital and postfrontal and containing the supratemporal fenestra; a distinct antorbital ridge on dorsal surface of prefrontal; postfrontal ridge forming medial border of ���supratemporal trough���; hypertrophied postorbital and squamosal carrying a dorsal ridge to form lateral border of the ���supratemporal trough���; temporal process of parietal short, roughly 1/ 3 of parietal length; a deep U-shaped post-temporal incision at the midline of the occiput; low-crowned dentition; strongly elongated iliac blade with parallel dorsal and ventral borders; presence of a distinct spike-like posterior process of ischium. Anatomical description The holotype (PKUP V2001) includes a nearly complete skull, mandibles and postcranial skeleton missing minor parts of the limbs (Fig. 2). The entire skeleton is preserved in a volcanic shale slab, and was exposed in dorsal view after preparation of the specimen. Skull roof. The cranial skeleton is nearly completely preserved, and is dorsoventrally compressed as in other choristoderes (Fig. 3). The general configuration of the skull in dorsal view, however, is sharply different from the type genus Monjurosuchus, as the skull of the new form is substantially more elongated, with the antorbital portion slightly longer than the postorbital portion. The total length of the skull is approximately 103 mm, with the antorbital portion 40 mm, the orbit 25 mm, and the postorbital portion 38 mm long. The external narial openings are paired, elongated and are nearly terminal in position. With a strong interorbital constriction, the orbit is large but more dorsally than laterally orientated. The orbit is longer than wide, differing from that in Monjurosuchus (Gao et al., 2000). The supratemporal fenestra of Philydrosaurus proseilus is unique: it is small and connects with an anterior trough formed by the postorbital and postfrontal (Fig. 3). The infratemporal fenestra is completely closed by expansion of the surrounding bones, including the postorbital, quadratojugal and squamosal. The closing of the infratemporal fenestra is apparently a synapomorphic feature of the Monjurosuchidae, as it also occurs in the type genus Monjurosuchus (Gao et al., 2000). No post-temporal fenestra is developed: the extremely short temporal process of the parietal is in full contact with the squamosal and the quadrate (see below). The premaxillae are small, paired elements that meet along a midline suture. The body of the premaxilla is penetrated by several foramina for nerves and vessels, and these are associated with small bumps developed as surface ornamentation. Each premaxilla has a slender but well-defined dorsal process that forms 1/3���1/2 of the medial border of the external nares (the dorsal process of the left premaxilla extends to the level of the posterior border of the narial openings). The posterior end of the dorsal process narrows between the slender anterior processes of the nasals that form the posterior part of the medial border of the external nares. The posterior margin of the premaxillary body is notched for the anterior border of the narial opening, and lateral to the opening is the irregular suture that marks the articulation between the premaxilla and maxilla. Posteroventrally, the premaxilla has a well-developed tooth-bearing process extending to the midlevel of the narial opening. The nasals are paired, very narrow, elongated (20 mm long) and extend posteriorly to contact the prefrontal at the midlevel of the snout. Anteriorly, the nasal forms the posterior, but not part of the lateral, border of the narial opening. A slender anterior process of the bone contacts the dorsal process of the premaxilla to form a large part of the internarial septum. Laterally, the nasal has a long, straight suture with the maxilla, and posteriorly it is developed as a slender process that pinches between the anterior process of the prefrontal and the dorsal rim of the maxilla. The prefrontals are paired and elongated, being the same length (20 mm) as the nasals. A midline sutural contact between the prefrontals is clearly visible as in other choristoderes generally; however, a narrow opening is evident within the midline suture; such a prefrontal fontanel has never previously been recorded in other choristoderes, including a small subadult skull of Champsosaurus (RTMP 94.163.1, see Gao & Fox, 1998) from the Upper Cretaceous Dinosaur Park Formation of western Canada. The dorsal surface of the prefrontal of this new animal is unique in having a sharp elongate ridge running anteriorly from the orbital rim to the level of nasal/prefrontal articulation. The prefrontal contacts the nasal anteriorly, the frontal posteriorly, and the lacrimal and maxilla laterally. The frontals are paired, elongated and hourglassshaped, with a strong interorbital constriction. Having a greatest length of 26 mm, the frontal is longer than the prefrontal, and equal in length to the parietal (26 mm). The anterior part of the frontal is broken bilaterally, but it seems to lack an anteromedial process. The anterolateral processes are in articulation with the prefrontal but barely contact the lacrimal. The anterior and posterior width of the paired frontals is 11 mm, and the interorbital width, 5 mm. The frontal contacts the postfrontal posterolaterally along a short straight suture, and meets the parietal posteriorly at a contact that is roughly transverse to the long axes of these bones. The latter suture is at about the same level as the anterior border of the ���supratemporal trough��� or the posterior border of the orbits. The parietals are paired, having a total length of 26 mm, including the short temporal process. The dorsal surface of the parietal table is smooth, but a prominent longitudinal ridge is developed that flanks the lateral border of the table. This lateral ridge also forms the medial border of the small supratemporal fenestra, and is continuous anteriorly with the ridge of the postfrontal that extends to the posteromedial border of the orbit. There is no parietal foramen. The temporal process of the parietal is extremely short (9 mm long), roughly 1/3 of the total length of the bone (26 mm). The two temporal processes form a deep Ushaped post-temporal incision medially on the occiput (Fig. 3). The maxillae are well preserved on both sides. As in other choristoderes, the maxilla lacks a well-defined dorsal process, but inrolls medially to contact the elongated nasal bone. The maxilla has a short anterior process that is in articulation with the premaxilla and forms the lateral border of the narial opening. Medially, the maxilla has a long suture with the nasal and posteriorly contacts the lacrimal: the lateral wall of the maxilla sharply descends to meet the lacrimal and jugal. The slender posterior process of the maxilla is largely obscured on the right side, but the betterexposed left side seems to show it terminates at a level close to the posterior border of the orbit. This interpretation is supported by the fact that the lower dentition also terminates at this level, as shown by the disarticulated left mandible (Fig. 3). Anterior to the orbit, the lacrimal is located in a longitudinal depression that is deep and wide in its posterior part but gradually becomes narrow and shallow anteriorly, terminating not far behind the narial opening. The lacrimal is quite narrow and elongated (25 mm long), with its anterior extremity reaching beyond the level of the nasal/prefrontal suture. Medially, it meets the prefrontal, and laterally the maxilla and anterior process of the jugal. The medial edge of the lacrimal contacts the prefrontal in the longitudinal depression developed dorsally on the snout (Fig. 3). The posterior border of the lacrimal is deeply notched for the anterior border of the orbit, and the dorsal rim of the orbit is ornamented with prominent tuberosities. The lacrimal foramen is identifiable on both sides, despite the dorsoventral compression of the skull. This foramen penetrates the lacrimal, the primitive condition differing from the derived pattern in which the foramen is bordered by the lacrimal, prefrontal and palatine as in neochoristoderes (see Gao & Fox, 1998). The postorbital and postfrontal are separate, and both enter the posterior border of the orbit (Fig. 3). This morphology is primitive in diapsids and differs from the fused condition in some but not all derived choristoderes (Gao & Fox, 1998; Evans & Manabe, 1999; Ksepka, Gao & Norell, 2005). In dorsal view, the postfrontal is substantially smaller than the postorbital, but overlaps the latter element in the ���supratemporal trough.��� Dorsally, the postfrontal displays a short but prominent ridge that runs from the posterior border of the orbit to meet the lateral ridge of the parietal. Medial to this ridge, the postfrontal contacts both the frontal and the parietal along a short, straight suture; the posterior extension of this suture enters obliquely into the ���supratemporal trough��� that separates the postfrontal from a small lateral projection of the parietal. Also within the trough, the postfrontal overlaps the postorbital to form much of the anterior border of the supratemporal fenestra. Lateral to the postfrontal, the postorbital is greatly hypertrophied as the main element that closes the infratemporal fenestra. Dorsally, the postfrontal displays a strongly developed ridge that joins with the ridge from the squamosal to form the lateral border of the ���supratemporal trough���. Lateral to the dorsal ridge, the postorbital has an extensive sutural contact posteriorly with the squamosal and ventrally with the quadratojugal. As a diagnostic feature of the family Monjurosuchidae, these three elements together have entirely closed the infratemporal fenestra, leaving no trace of the opening. The jugal and the quadratojugal are preserved on both sides in the holotype. The anterior process of the jugal forms the lower border of the orbit and extends anteriorly to the lateral surface of the posterior process of the maxilla. The anterior tip of the process is damaged on both sides, but it probably terminated at a level slightly anterior to the orbit, as interpreted from the articular surface of the jugal on the maxilla. The postorbital, or dorsal, process of the jugal is low but well defined, forming a small part of the posteroventral border of the orbit. From the postorbital process, the dorsal margin of the jugal descends posteriorly to meet the quadratojugal. The short posterior process of the jugal fits into a small notch on the quadratojugal slightly anterior to the craniomandibular joint. The quadratojugal is a much smaller bone in comparison to the jugal and squamosal. It is located between the jugal and the squamosal, and dorsally contacts the enlarged postorbital. The squamosal is greatly expanded as a major element that closes the infratemporal fenestra. The dorsolateral exposure of the squamosal is roughly a triangular plate, with a flattened surface rugosely ornamented. In dorsal view, the medial extension of the squamosal forms part of the floor of the ���supratemporal trough���, and meets the temporal process of the parietal. The anterior border of this medial extension is thin, slightly concave and overlaps the quadrate. Posteriorly, this extension has two prominent processes separated from one another by a small notch (Fig. 3). The medial process forms a small part of the lateral border of the narrow U-shaped post-temporal incision, and the lateral process forms the posterior extremity of the temporal region. Braincase and palate: Only a small part of the lateral wall of the braincase is exposed in the supratemporal fenestra within the ���supratemporal trough���, and the lateral wall is badly crushed on both sides. However, it is clear that a downgrowth of the parietal forms a small part of the braincase wall, and it has a dorsally concave border for articulation with a small bone, presumably the neomorph as in other known choristoderes (Fox, 1968). A neomorph occurs in neochoristoderes and possibly in Cteniogenys (Evans, 1990), but this element cannot be identified positively for this new form because of breakage and distortion of the braincase. By the same token, a pterygoquadrate foramen is not identifiable on the new specimen, and the ventral parts of the braincase cannot be exposed without damaging the specimen; therefore, knowledge of the morphology of this part of the skull must await discovery of better preserved material. Within the orbit, the pterygoid and ectopterygoid are partly exposed in dorsal view. The short medial process of the ectopterygoid forms the posterior border of the suborbital fenestra and overlaps the short lateral process of the pterygoid. A slender anterior process of the ectopterygoid extends along the medial side of the anterior process of the jugal, terminating slightly anterior to the midlevel of the suborbital fenestra; therefore, the short anterior extension of the process allows a small part of the jugal to enter the lateral border of the suborbital fenestra (Fig. 3). The anterior extension of the pterygoid is greatly widened as a plate that forms the entire medial border of the suborbital fenestra. The midline suture of the pterygoids is obscured by the frontal, and thus, the taxonomically significant feature of the interpterygoid foramen cannot be observed in this specimen. The anterior end of the pterygoid is badly damaged on both sides, and the pattern of the pterygoid/palatine articulation cannot be determined. Posterior to the suborbital fenestra, only a small part of the subtemporal fenestra is exposed behind the medial process of the ectopterygoid; therefore, the actual size of the fenestra is unknown in the holotype of the new taxon. Disarticulated from the palate, the left palatine is exposed in close association with the skull (partly obscured by the left maxilla). Most of the palatine is a thin plate, but it is slightly thickened posteriorly at the notch for the suborbital fenestra. Along the dorsomedial border of the palatine, the articular surface for the greatly elongated vomer can be seen. This surface is about half the width of the palatine, indicating an extensive articulation between the two bones. Although exposed in dorsal view, the exposed lateral edge seems to show that the palatine has a toothed ventral surface, but the distribution pattern of the palatal teeth cannot be ascertained in the absence of a ventral view. Also disarticulated from the skull, the right vomer is nearly completely preserved in close association with a hyoid bone. Exposed in dorsal view, the vomer is greatly elongated, with a total length of 50 mm. The anterior extremity of the vomer is narrowly pointed but deeply bifurcated by a fissure 10 mm in length. About 19 mm from the anterior tip, a small vomerine foramen opens close to the medial border of the bone. Along this border, a well-defined ridge runs from the vomerine foramen to the posterior end of the anterior fissure. The vomer posteriorly has a notch for articulation with the pterygoid and posterolaterally an articular surface for the palatine. Lying to the side of the vomer is a slender bone 32 mm in length. This is probably one of the hyoid elements that are also known for some champsosaurs (Gao & Fox, 1998: fig. 5). Mandibles: Before burial, both mandibles of the holotype (PKUP V2001) were disarticulated from the skull (Fig. 3). The left mandible is nearly completely preserved and is exposed in medial view; the dentary of the right mandible is exposed in lateral view but the postdentary parts are missing. The total length of the mandible is 83 mm as measured from the nearly complete left element. With a total length of 63 mm, the dentary is the longest mandibular bone and covers a large part of the lateral side of the lower jaw. As shown in the right mandible, the strongly convex lateral surface of the dentary is penetrated by rows of small foramina. The foramina on the anterior part of the jaw are closely spaced, while those on the posterior part are increasingly widely spaced; they are accompanied by narrow and posteriorly extending grooves that housed lateral cutaneous branches of the mandibular ramus of the trigeminal nerve and accompanying blood vessels. The posterior end of the dentary is bifurcated for articulation with the surangular bone (missing on the right mandible along with the other postdentary elements). Medially, the dentary carries a well-developed subdental shelf, the depth of which is roughly equal to the tooth height. Below the shelf is the Meckelian canal that opens medially as a narrow groove for the anterior one-third of the dentary length, but is closed by the splenial for the posterior twothirds. Anteriorly, the mandibular symphysis is extremely short and lacks a posterior extension as seen in advanced neochoristoderes. The splenial has been crushed into the Meckelian canal, but it shows that its posterior extremity terminates as a short tongue slightly beyond the posterior end of the tooth row. This tongue is wedged between the coronoid and the anterior extension of the prearticular, but may have a short ventral sutural articulation with the angular bone. The coronoid is a small triangular bone without a prominent dorsal process. It sends a short posteroventral process to articulate with the splenial and the prearticular, thereby contributing to a small part of the lower border of the mandibular fossa. Behind the fossa is the articular surface for the craniomandibular joint. From the length of the mandible, this joint is interpreted as being located roughly at the same level as the occipital condyle. The articular surface is at the posterior extremity of the jaw, and thus a retroarticular process is undeveloped in this animal. Dentition: On the holotype specimen, marginal teeth are observed on both upper and lower jaws. Anteriorly, the upper dentition is exposed on both sides, but its more posterior parts are imbedded in matrix making the total number of the maxillary teeth indeterminable. As a result of the disarticulation of the lower jaws from the skull, the lower dentition on the left is exposed in medial view and that on the right in lateral view (Fig. 3). The left dentary has as many as 41 teeth preserved, with, Published as part of Gao, Ke-Qin & Fox, Richard C., 2005, A new choristodere (Reptilia: Diapsida) from the Lower Cretaceous of western Liaoning Province, China, and phylogenetic relationships of Monjurosuchidae, pp. 427-444 in Zoological Journal of the Linnean Society 145 (3) on pages 428-437, DOI: 10.1111/j.1096-3642.2005.00191.x, http://zenodo.org/record/4687363, {"references":["Gao K, Evans S, Ji Q, Norell M, Ji S. 2000. Exceptional fossil material of a semi-aquatic reptile from China: the resolution of an enigma. Journal of Vertebrate Paleontology 20: 417 - 421.","Gao KQ, Fox RC. 1998. New choristoderes (Reptilia: Diapsida) from the Upper Cretaceous and Palaeocene, Alberta and Saskatchewan, Canada, and phylogenetic relationships of Choristodera. Zoological Journal of the Linnean Society 124: 303 - 353.","Evans SE, Manabe M. 1999. A choristoderan reptile from the Lower Cretaceous of Japan. Special Papers in Palaeontology 60: 101 - 119.","Ksepka DT, Gao K-Q, Norell MA. 2005. A new choristodere from the Cretaceous of Mongolia. American Museum Novitates 3468: 1 - 22.","Fox RC. 1968. Studies of Late Cretaceous vertebrates I: The braincase of Champsosaurus Cope (Reptilia- Eosuchia). Copeia 1: 100 - 109.","Evans SE. 1990. The skull of Cteniogenys, a choristodere (Reptilia: Archosauromorpha) from the Middle Jurassic of Oxforshire. Zoological Journal of the Linnean Society 99: 205 - 237.","Evans SE. 1988. The early history and relationships of the Diapsida. In: Benton MJ, ed. The phylogeny and classification of the tetrapods 1: Amphibians, reptiles, birds. Oxford: Clarendon Press, 221 - 260.","de Braga M, Rieppel O. 1997. Reptile phylogeny and the interrelationships of turtles. Zoological Journal of the Linnean Society 120: 281 - 354.","Sigogneau-Russell D. 1981. Etude osteologique du reptile Simoedosaurus (Chostiodera). Iie partie: squelette postcranien. Annales de Paleontologie Vertebres 67: 61 - 140.","Gauthier JA, Kluge AG, Rowe T. 1988. The early evolution of the Amniota. In: Benton MJ, ed. The phylogeny and classification of the tetrapods 1: Amphibians, reptiles, birds. Oxford: Clarendon Press, 103 - 155.","Rieppel O. 1993. Euryapsid relationships: a preliminary analysis. Neues Jahrbuch fur Geologie und Palaontologie 188: 241 - 264.","Romer AS. 1956. Osteology of the reptiles. Chicago / London: The University of Chicago Press.","Reisz RR. 1981. A diapsid reptile from the Pennsylvanian of Kansas. Special Publication of the Museum of Natural History, University of Kansas 7: 1 - 74.","Rieppel O. 1994. Osteology of Simosaurus gaillardoti, and the phylogenetic interrelationships of stem-group Sauropterygia. Fieldiana (Geology) N S 28: 1 - 85.","Carroll RL, Currie PJ. 1991. The early radiation of diapsid reptiles. In: Schultze H-P, Trueb L, eds. Origins of the higher groups of tetrapods. Ithaca / London: Comstock Publishing, 354 - 424."]}

GENUS PHILYDROSAURUS GEN. NOV. Type species: Philydrosaurus proseilus sp. nov. Diagnosis: As for the type and only known species. Range: Early Cretaceous, western Liaoning, China., Published as part of Gao, Ke-Qin & Fox, Richard C., 2005, A new choristodere (Reptilia: Diapsida) from the Lower Cretaceous of western Liaoning Province, China, and phylogenetic relationships of Monjurosuchidae, pp. 427-444 in Zoological Journal of the Linnean Society 145 (3) on page 428, DOI: 10.1111/j.1096-3642.2005.00191.x, http://zenodo.org/record/4687363

Suborder Neochoristodera Consisting of the traditional familial groups, Champsosauridae and Simoedosauridae (Fig. 6), the taxon Neochoristodera as named by Evans & Hecht (1993) designates the same clade as the name Champsosauriformes Hay, 1929. However, the name Neochoristodera has been widely accepted, while the name Champsosauriformes has not been mentioned in the literature for more than 50 years; thus, the latter name is here treated as a nomen oblitum (a forgotten name). The suborder Neochoristodera is a robust clade that can be unambiguously diagnosed by the following synapomorphies: external nares terminal and confluent (1-2; possibly also in Cteniogenys); dorsal flange of maxilla low and strongly inrolled (6-2); nasals elongate and fused (9-2); nasals intervene between premaxillae (10-1); orbit small and dorsally directed (11- 2); dorsal process of squamosal slender and elongate to middle level of infratemporal fenestra (13-1); supratemporal fenestra posteriorly flared and substantially larger than orbit (14-2); vomer/maxilla contact present (19-1); palatal foramen opens between pterygoid and palatine without contribution from the vomer (21-2); quadratojugal bears a cotyle meeting a rounded quadrate articulation (26-1); pterygoid process of quadrate low, slender and horizontal with a reduced pterygoid facet (27-1); nasopalatal trough present (29-1); pterygoids meeting in a long midline suture (31-1); interpterygoid vacuity small and near posterior border of suborbital fenestra (32-1); basal infolding of tooth enamel present (37-1); pterygoquadrate foramen present and enclosed between neomorph and quadrate (41-1); neurocentral sutures remain open in adult (51-1); presacral vertebral centra short and spool-like (53-1); number of ossified carpal elements seven or fewer (60-1); bone structure pachyostotic in adult (62-2); iliac blade dorsal margin essentially horizontal, blade expanded (63-1); clavicular facet on interclavicle continuous across midline in smooth curve (66-1); constriction of ilium between iliac blade and acetabulum as a well-defined neck (74-1; reversal in Ikechosaurus).

MONJUROSUCHIDAE AS A MONOPHYLETIC FAMILY Two genera and species are currently classified in the family Monjurosuchidae: the type species Monjurosuchus splendens from the Yixian Formation exposed near Lingyuan, and Philydrosaurus proseilus gen. et sp. nov. described here from the Chiufotang Formation exposed near Chaoyang. With inclusion of these two taxa in a revised data set (Appendix 2), a parsimony analysis revealed that the monophyly of the family is supported by the following synapomorphies: dorsal process of squamosal absent (13-2; unknown for Hyphalosaurus and Shokawa); supratemporal fenestra smaller than orbit (14-0; character reversal); infratemporal fenestra closed (15-1; homoplastic in Lazarussuchus, see Evans & Klembara, 2005); post-temporal fenestra absent (47-1; also in Simoedosauridae); posterior process of ischium strongly developed (75-1; uniquely diagnostic for the family Monjurosuchidae). A POSSIBLE MONJUROSUCHIDAE ���HYPHALOSAURIDAE CLADE Hyphalosaurus and Shokawa are two morphologically peculiar, long-necked choristoderes known from the Lower Cretaceous of China and Japan (Evans & Manabe, 1999; Gao, Tang & Wang, 1999). Both MPTs (Fig. 6) recognize a Hyphalosaurus ��� Shokawa clade, for which the name Hyphalosauridae (fam. nov.) is proposed here. The grouping of the two forms in the family Hyphalosauridae is unambiguously supported by two uniquely derived character states of the two genera of the family: cervical vertebral count more than 16 (67-1); and caudal neural spines tall and narrow (72-1). Because the skull morphology is either poorly known or entirely unknown for the two forms, the diagnosis of the family would be enhanced by further study of the known material and discovery of new material. Within the phylogenetic framework of the Choristodera, both Monjurosuchidae and Hyphalosauridae are recognized as robust familial groups outside the neochoristoderan clade; however, the two MPTs differ from one another in placing the two families as either successive sister groups with the Neochoristodera (Fig. 6A), or grouped together as the sister group with the Neochoristodera (Fig. 6B). The grouping of Monjurosuchidae with Hyphalosauridae as sister groups is weakly supported by the following two character states: lateral borders of frontals strongly constricted (3-1; unknown for Shokawa); and orbit large and dorsally orientated (11-1; unknown for Shokawa). Because the current data matrix contains a large amount of missing data for the basic taxa of the two families, the robustness of the possible Monjurosuchidae��� Hyphalosauridae clade must be tested further when new data become available., Published as part of Gao, Ke-Qin & Fox, Richard C., 2005, A new choristodere (Reptilia: Diapsida) from the Lower Cretaceous of western Liaoning Province, China, and phylogenetic relationships of Monjurosuchidae, pp. 427-444 in Zoological Journal of the Linnean Society 145 (3) on page 438, DOI: 10.1111/j.1096-3642.2005.00191.x, http://zenodo.org/record/4687363, {"references":["Evans SE, Klembara J. 2005. A choristoderan reptile (Reptilia: Diapsida) from the lower Miocene of northwest Bohemia. Journal of Vertebrate Palaeontology 25: 171 - 184.","Evans SE, Manabe M. 1999. A choristoderan reptile from the Lower Cretaceous of Japan. Special Papers in Palaeontology 60: 101 - 119.","Gao KQ, Tang Z, Wang X. 1999. A long-necked diapsid reptile from the Upper Jurassic / Lower Cretaceous of Liaoning Province, northeastern China. Vertebrata Palasiatica 37: 1 - 8."]}

Heishanlestes, new genus TYPE SPECIES: Heishanlestes changi, n.sp. DIAGNOSIS: As for the type and only species. ETYMOLOGY: ‘‘ Heishan’ ’ is the name of the county in northeastern China in which the materials of the type species were collected. DISTRIBUTION AND AGE: As for the type and only species.

Heishanlestes changi, new species Figures 1���5, table 1 HOLOTYPE: IVPP V 7480, a nearly complete right dentary containing p1���4, m1���6 (figs. 1, 3, 4, 5). REFERRED SPECIMENS: IVPP V 7481, anterior part of a left dentary containing an erupting canine, p1���4, m1 (figs. 2, 3); IVPP V 7482, anterior part of a left dentary containing p4, m1, and alveolus for p3 (fig. 2). ETYMOLOGY: ������changi������ is for Professor Zhenglu Chang, from Liaoning Technical University, for his important contribution to the study of Mesozoic stratigraphy and paleontology of northeastern China during the past 50 years. DIAGNOSIS: Dental formula: ci, p1���4, m1��� 6, differing from Spalacotherium tricuspidens and Spalacolestes cretulablatta, which have seven lower molars; premolars tightly spaced and overlapping; m1 with obtuse trigonid angle; m5���6 reduced in size and with a subcylindrical cusp arising from the center of the trigonid; differing from Zhangheotherium and Maotherium in having a strong pterygoid crest but absence of an enlarged first incisor, absence of a Meckelian groove, and reduction of cingular cusps on lower molars. LOCALITY: Badaohao, Heishan County, Liaoning Province, China (coordinates: 418509N and 1218559E). GEOLOGIC OCCURRENCE: Shahai Formation. AGE: Aptian, Early Cretaceous., Published as part of HU, YAO-MING, FOX, RICHARD C., WANG, YUAN-QING & LI, CHUAN-KUI, 2005, A New Spalacotheriid Symmetrodont from the Early Cretaceous of Northeastern China, pp. 1-20 in American Museum Novitates 3475 (1) on page 4, DOI: 10.1206/0003-0082(2005)475[0001:ANSSFT]2.0.CO;2, http://zenodo.org/record/4712609

HU, YAO-MING, FOX, RICHARD C., WANG, YUAN-QING, LI, CHUAN-KUI (2005): A New Spalacotheriid Symmetrodont from the Early Cretaceous of Northeastern China. American Museum Novitates 3475 (1): 1-20, DOI: 10.1206/0003-0082(2005)475[0001:ANSSFT]2.0.CO;2, URL: http://www.bioone.org/perlserv/?request=get-abstract&doi=10.1206%2F0003-0082(2005)475%5B0001%3AANSSFT%5D2.0.CO%3B2