Cretaceous Research xxx (2013) 1e10 Contents lists available at SciVerse ScienceDirect Cretaceous Research journal homepage: www.elsevier.com/locate/CretRes Kazaklambia convincens comb. nov., a primitive juvenile lambeosaurine from the Santonian of Kazakhstan Phil R. Bell a, *, Kirstin S. Brink b a b Pipestone Creek Dinosaur Initiative, 10001 84th Avenue, Clairmont, Alberta T0H 0W0, Canada Department of Ecology and Evolutionary Biology, University of Toronto Mississauga, 3359 Mississauga Road, Mississauga, Ontario L5L 1C6, Canada a r t i c l e i n f o a b s t r a c t Article history: Received 11 October 2012 Accepted in revised form 18 May 2013 Available online xxx The holotype of ‘Procheneosaurus’ convincens, a juvenile lambeosaurine, is the most complete hadrosaurid known from Kazakhstan. North American species of Procheneosaurus are considered juveniles of Corythosaurus, Hypacrosaurus, and Lambeosaurus, rendering the generic name unusable. A replacement name, Kazaklambia convincens comb. nov., is herein proposed as this specimen is morphologically distinct from other Eurasian taxa and known juvenile lambeosaurines at a similar ontogenetic stage in having a prefrontal process of the postorbital with a dorsal thickening forming a dome lateral to the frontal dome, doming of the nasal anterodorsal to the orbit, and a frontal length/width ratio <1. The juvenile status of Kazaklambia makes phylogenetic placement difficult; however, morphometric and morphological information (particularly in relation to the hollow cranial crest and the length of the frontal) suggest a close affiliation with the basal lambeosaurines Amurosaurus and Tsintaosaurus, and support the hypothesis for an Asian origin for Lambeosaurinae. Ó 2013 Elsevier Ltd. All rights reserved. Keywords: Lambeosaurinae Asia Taxonomy Cretaceous 1. Introduction Lambeosaurine hadrosaurid dinosaurs are a diverse and wellknown group from the Late Cretaceous of Laurasia (Horner et al., 2004). Their fossil record in North America includes nearly complete ontogenetic series, composed of embryonic, nestling, juvenile, subadult, and adult specimens (Horner and Currie, 1994; Evans et al., 2005, 2007; Brink et al., 2011). Growth stages preceding the adult growth stage lack full expression of the cranial crests that characterize this group. Consequently, prior to the work of Dodson (1975), specimens without well-developed cranial crests were classified into a distinct genus with three species, Procheneosaurus (¼Tetragonosaurus) erectofrons, P. praeceps and P. cranibrevis (Lull and Wright, 1942; Sternberg, 1953; Evans et al., 2005). These species are now identifiable to the genus level as juveniles of Corythosaurus spp. and Lambeosaurus spp. based on morphometric data and discrete characters (Dodson, 1975; Evans et al., 2005). A specimen initially identified as P. erectofrons (AMNH 5461) is now considered as a juvenile of Hypacrosaurus stebingeri from the Two Medicine Formation of Montana, USA (Horner and Currie, 1994). Another juvenile taxon, Cheneosaurus tolmanensis, is now * Corresponding author. E-mail addresses: philbyb@gmail.com (P.R. Bell), kirstin.brink@mail.utoronto.ca (K.S. Brink). considered as Hypacrosaurus altispinus (Evans, 2010). Juveniles and subadults of Parasaurolophus are also identifiable to the genus level based on the anatomy of the skull roof and incipient cranial crest (Evans et al., 2007, 2009). In 1961, a virtually complete skull and skeleton (PIN 2230/1) of a lambeosaurine hadrosaurid was discovered in rocks of the Dabrazinskaya Svita at the SyukeSyuk well site, 45 km northwest of Tashkent (Uzbekistan) in southeastern Kazakhstan (Fig. 1). The age of the Dabrazinskaya Svita is poorly constrained; however, it is generally regarded as Santonian in age (Rozhdestvensky, 1968, 1974; Averianov and Nessov, 1995). The skeleton was largely intact and articulated except for the right pes, both mani, and most of the preorbital region of the skull, which was lost to erosion prior to discovery. Rozhdestvensky (1968) named the new animal Procheneosaurus convincens based on the low, hollow nasal crest, which was similar to the North American ‘procheneosaurs’ from the Belly River Group of Alberta. Weishampel and Horner (1990) and Horner et al. (2004) synonymized P. convincens with Jaxartosaurus aralensis but did not provide any justification for this reassignment. Norman and Sues (2000) argued on the grounds of stratigraphic separation and the taxonomic differences described by Rozhdestvensky (1968) that it was probably prudent to retain P. convincens as a distinct taxon, but considered its status as questionable. Following  ska and Osmólska (1981a, b), Norman and Sues (2000), Maryan suggested that a replacement generic name might be necessary for P. convincens. 0195-6671/$ e see front matter Ó 2013 Elsevier Ltd. All rights reserved. http://dx.doi.org/10.1016/j.cretres.2013.05.003 Please cite this article in press as: Bell, P.R., Brink, K.S., Kazaklambia convincens comb. nov., a primitive juvenile lambeosaurine from the Santonian of Kazakhstan, Cretaceous Research (2013), http://dx.doi.org/10.1016/j.cretres.2013.05.003 2 P.R. Bell, K.S. Brink / Cretaceous Research xxx (2013) 1e10 Fig. 1. Locality map of Kazakhstan in eastern Europe showing the type locality of Kazaklambia convincens comb. nov. (star). Four other hadrosauroids are known from Kazakhstan: Aralosaurus tuberiferus, Arstanosaurus akkurganensis, Batyrosaurus rozhdestvenskyi, and Jaxartosaurus aralensis (Rozhdestvensky, 1968; Norman and Sues (2000), Godefroit et al., 2004b, 2012). Arstanosaurus akkurganensis and Batyrosaurus rozhdestvenskyi both come from the Aggurgan locality in Central Kazakhstan. Norman and Sues (2000) considered the partial maxilla and distal femur that comprise the holotype material of A. akkurganensis a nomen dubium and we follow their reasoning here. Batyrosaurus is described as the youngest non-hadrosaurid hadrosauroid (Godefroit et al., 2012). Both Aralosaurus and Jaxartosaurus are recovered in recent phylogenetic analyses as basal lambeosaurines (Prieto-Marquez, 2010a; Evans, 2010; Godefroit et al., 2004a,b; Godefroit et al., 2008) or as the sister-taxon to Hadrosauridae (Sues and Averianov, 2009). Aralosaurus tuberiferus is from the (?) Turonian SakheSakh locality in central Kazakhstan, and Jaxartosaurus aralensis is from the same region as P. convincens but from a slightly lower (Santonian) stratigraphic level. Given that Procheneosaurus is no longer a valid genus based on the synonymy of North American taxa (Dodson, 1975) and previous claims that P. convicens may be a juvenile of Jaxartosaurus (Weishampel and Horner, 1990; Horner et al., 2004), the holotype of P. convicens is in need of revision. Considering that specimens previously assigned to Procheneosaurus from North America are now identifiable to the genus and some to the species level based on the morphology of the skull, we focus our redescription on the holotype skull of P. convincens. Comparisons differentiate P. convicens from lambeosaurines of the same ontogenetic stage and from the Kazakstanian lambeosaurines Jaxartosaurus and Aralosaurus. A morphometric analysis is performed to compare PIN 2230/1 to Laurasian hadrosauroids known from a variety of growth stages. Institutional abbreviationsdAMNH, American Museum of Natural History, New York, USA; PIN, Palaeontologiceski Institut, Academii Nauk, Moscow, Russia. 2. Systematic palaeontology Dinosauria Owen, 1842 Ornithischia Seeley, 1887. Ornithopoda Marsh, 1881 Hadrosauridae Cope, 1869 Lambeosaurinae Parks, 1923 Kazaklambia, gen. nov. Type Species. Kazaklambia convincens (Rozhdestvensky, 1968) Diagnosis. As for type and only species. Etymology. Prefix references Kazakhstan, the country in which the holotype specimen was found; lambia denotes the affiliation to Lambeosaurinae. Kazaklambia convincens (Rozhdestvensky, 1968) comb. nov., Figs. 2 and 3 Procheneosaurus convincens Rozhdestvensky, 1968 Etymology. Latinization of ‘convince’, referring to Rozhdestvensky’s (1968) conviction that this specimen proved a Cretaceous age for the Dabrazinskaya Svita. Emended diagnosis. At the juvenile ontogenetic stage, autapomorphies include a prefrontal process of postorbital with dorsal thickening forming dome lateral to frontal dome, doming of nasal anterodorsal to orbit, and a frontal length/width ratio <1. Holotype. PIN 2230/1, virtually complete skull and skeleton of an immature individual lacking most of the preorbital region of the skull, distal parts of the forelimbs, distal left hindlimb, and distal caudal vertebrae. Type locality. SyukeSyuk well site, southeastern Uzbekistan. Dabrazinskaya Svita (Santonian). 2.1. Description 2.1.1. Dermatocranium 2.1.1.1. Premaxilla. Only the posterior-most portion of the lateral process of the premaxilla is preserved where it contacts the lacrimal posteroventrally, prefrontal posterodorsally, and nasal dorsally. The suture between the premaxilla and the lacrimale prefrontal is nearly straight. The premaxilla is squared-off where it contacts the nasal, forming a right angle between the nasal and prefrontal contacts (Fig. 2A, B) similar to juvenile Corythosaurus (Evans et al., 2005) but unlike Hypacrosaurus, where the equivalent portion of the premaxilla is sharply attenuating (Evans, 2010). The anterior margin of the lateral premaxillary process, although damaged, is broadly v-shaped as in juvenile Corythosaurus (Evans Please cite this article in press as: Bell, P.R., Brink, K.S., Kazaklambia convincens comb. nov., a primitive juvenile lambeosaurine from the Santonian of Kazakhstan, Cretaceous Research (2013), http://dx.doi.org/10.1016/j.cretres.2013.05.003 Fig. 2. Holotype skull of Kazaklambia convincens comb. nov. (PIN 2230/1) in A. left lateral, C., posterior; and E. dorsal views. B, D, and F, interpretive illustrations of the same. Arrowhead indicates dorsal extent of jugal. Broken surfaces denoted by cross-hatching. Light grey regions are invertebrate borings; dark grey denotes supporting armature for mount. Abbreviations: An, angular; Ar, articular; Bo, basioccipital; Ex, exoccipital; D, dentary; Fm, foramen magnum; Fr, frontal; J, jugal; La, lacrimal; Mx, maxilla; Na, nasal; Pa, parietal; Pfr, prefrontal; Pmxl, lateral process of the premaxilla; Po, postorbital; Pod, postorbital dome; Pt, pterygoid; Q, quadrate; Soc, supraoccipital; Sq, squamosal; Su, surangular. Please cite this article in press as: Bell, P.R., Brink, K.S., Kazaklambia convincens comb. nov., a primitive juvenile lambeosaurine from the Santonian of Kazakhstan, Cretaceous Research (2013), http://dx.doi.org/10.1016/j.cretres.2013.05.003 4 P.R. Bell, K.S. Brink / Cretaceous Research xxx (2013) 1e10 et al., 2005), and therefore strongly suggestive of the presence of a premaxillaryenasal fontanelle. The premaxillaryenasal fontanelle is present in juvenile Corythosaurus, Lambeosaurus, Hypacrosaurus stebingeri, and Velafrons coahuilensis, but is closed in all specimens  ska and Osmólska, 1979; Evans, 2010; Brink of H. altispinus (Maryan et al., 2011). 2.1.1.2. Maxilla. Only the posterior half of the maxilla is preserved and very little of its general morphology can be observed due to the surrounding elements and articulated lower jaws. However, there is no indication that they differed from the general lambeosaurine beauplan. 2.1.1.3. Nasal. Together, the paired nasals form a hollow, low, rounded crest anterodorsal to the orbit that is typical of most juvenile lambeosaurines (Fig. 2A, B). The apex of the crest is higher than the apex of the frontal dome, and comparatively more developed than juveniles of Corythosaurus and Lambeosaurus of a similar size, although this feature is ontogenetically variable (Evans et al., 2005; Evans, 2010). The anterior part of the nasal is missing; therefore, the forward extent of the crest cannot be determined. Consequently, it is unknown whether Kazaklambia possessed anteriorly bifurcating nasals as in Hypacrosaurus and Corythosaurus or unbranched as in Lambeosaurus (Evans et al., 2005; Brink et al., 2011) and possibly Velafrons (Gates et al., 2007). The nasals meet along a straight midline suture that becomes indistinct anteriorly, which is probably a preservational artefact rather than due to fusion. In dorsal view, the posterior margin of the nasal is semicircular where it forms an extensive lap joint with the underlying frontal (Fig. 2E, F). A prominent invagination is present between the nasals at the nasalefrontal junction, which is occupied by a reciprocal spur of the combined frontals as in Jaxartosaurus and Amurosaurus (Godefroit et al., 2004a,b). The lateral margin of the nasal is constricted at about midlength by the prefrontal. A short contact with the postorbital is formed at the posterolateral corner of the nasal between the frontal and prefrontal contacts. 2.1.1.4. Lacrimal. The lacrimal is a roughly trapezoidal element that forms part of the anterior margin of the orbit. It contacts the jugal ventrolaterally and maxilla ventrally. Dorsally and anteriorly, it meets the premaxilla along a contact that is Z-shaped in lateral view (Fig. 2A, B). A superficially similar Z-shaped contact with the jugal is due to the broken edges of the anterior process of the jugal. Although broken anteriorly, the lacrimal is longer anteroposteriorly than it is high. 2.1.1.5. Prefrontal. The prefrontal is a platy element that forms the anterodorsal margin of the orbit. The prefrontal is thickest medially where it also forms an anteroposteriorly elongate contact with the nasal. Posteriorly, the prefrontal attenuates between the frontal posteromedially and the postorbital posterolaterally. The postorbital and frontal contacts are almost at right angles when viewed from above (Fig. 2E, F). In lateral view, the prefrontal is arched to accommodate the curvature of the eye. Anteriorly, the prefrontal  ska and meets the premaxilla and lacrimal more ventrally. Maryan Osmólska (1979) commented on the presence of a supraorbital element that occupies this position in Kazaklambia. The presumed prefrontalesupraorbital suture is visible on both left and right sides of the skull extending anterolaterally in dorsal aspect (Fig. 2E, F); however, we cannot discount the possibility that these are in fact cracks. If the surpraorbital is indeed present, then it represents the only occurrence of this element in Lambeosaurinae. 2.1.1.6. Postorbital. In lateral aspect, the postorbital is roughly Tshaped. The jugal process descends at right angle to the main part of the anterior and posterior rami. Together, the jugal process and the anterior process form the posterodorsal margin of the orbit. Viewed laterally, the part of the orbit formed by the postorbital is arcuate, contrasting with the more abrupt right angle formed between the posterior ramus and the jugal process. The anterior ramus contacts the prefrontal anteriorly and frontal medially. On its dorsal surface, the anterior ramus is thickened to form a conspicuous dome (Fig. 2B). This dome is present on both left and right sides, which negates the possibility of it being a pathological or taphonomic feature. This dome is likely ontogenetic, as it is also present in immature specimens of Charonosaurus (Godefroit et al., 2001). A superficially similar structure is present in the largest Saurolophus angustirostris specimens (‘postorbital boss’ of Bell, 2011); however, in S. angustirostris, it is ornamented by a series of furrows and ridges. The roughly cylindrical posterior process forms an elongate scarf joint with the squamosal along its dorsolateral and medial surfaces. Due to the scarf joint, the posterior process appears forked in dorsal view, similar to H. stebingeri, Corythosaurus spp., and Lambeosaurus spp., but not H. altispinus. The corresponding region in Aralosaurus is broken and reconstructed. 2.1.1.7. Squamosal. The squamosal is a complex element forming the posterolateral corner of the skull (Fig. 2). Forming the lateral margin of the supraorbital fenestra, the anterior process meets the postorbital by way of a scarf join that extends the entire length of that process. A robust prequadratic process extends ventrally along part of the anterior edge of the quadrate. Immediately posterior to the prequadratic process, the squamosal forms a cotylus that encloses the head of the quadrate. A triangular postquadratic process extends posteroventrally and laterally; the posterior surface of which is met by the paroccipital process of the exoccipitaleopisthotic. The medial process is convex and rises dorsomedially above the supraoccipital to meet its counterpart at the skull midline (Fig. 2C, D). This process is similarly dorsomedially oriented in all other lambeosaurines except Jaxartosaurus in which the squamosals are flat lying (Godefroit et al., 2004b; Evans, 2010). There is no medial separation of the squamosals by the parietal as there is in Jaxartosaurus, Aralosaurus, Velafrons, some specimens of Hypacrosaurus, and Hadrosaurinae (Brink, 2009; Evans, 2010; Gates et al., 2011). 2.1.1.8. Quadrate. The dorsoventrally elongate quadrate is bowed posteriorly in lateral view as is characteristic of Lambeosaurinae (Horner et al., 2004). The dorsal quadrate head is enclosed anteriorly, dorsally, and posteriorly by the corresponding cotylus in the squamosal. Ventrally, it is mediolaterally expanded to form the articulating surface with the corresponding glenoid of the lower jaw. The articular surface consists of a lateral and smaller medial condyle. The medial condyle is situated dorsal to the lateral condyle and the two are separated by a shallow sulcus. Anteriorly, the quadratojugal contacts the quadrate by way of a lapped joint. Although the quadratojugal extends dorsally above the midheight of the quadrate, the semicircular quadratojugal facet (of the quadrate) is located ventral to the midheight of that element. The pterygoid flange is incompletely visible, extending anteromedially and broadly lapping the pterygoid along its posteromedial surface. A prominent dorsoventral groove separates the pterygoid process from the main part of the quadrate. 2.1.1.9. Jugal. Although it lacks the anterior-most part of the anterior process, the jugal is overall similar to juvenile corythosaurs such as Corythosaurus and Lambeosaurus (Evans, 2010). The anterior process is dorsoventrally expanded to the same height as the posterior process. On its medial surface, the anterior process broadly contacts the lacrimal dorsally and maxilla ventrally. The Please cite this article in press as: Bell, P.R., Brink, K.S., Kazaklambia convincens comb. nov., a primitive juvenile lambeosaurine from the Santonian of Kazakhstan, Cretaceous Research (2013), http://dx.doi.org/10.1016/j.cretres.2013.05.003 P.R. Bell, K.S. Brink / Cretaceous Research xxx (2013) 1e10 postorbital process is nearly straight and perpendicular to a line drawn between the ventral edges of the jugal flange and the anterior process. The distal half of the postorbital process extends posteromedially to the corresponding jugal process of the postorbital and terminates almost in line with the dorsal limit of the infratemporal fenestra (Fig. 2B). The jugal flange is prominent, effected by deep constrictions of the ventral margin of the posterior process and the main body of the jugal. The posterior process is hatchet-shaped and not angled as in H. altispinus (Evans, 2010). Although the quadratojugal is apparently present, no clear distinction between the posterior jugal and the quadratojugal can be made. This is without doubt a preservational feature rather than the true anatomical condition. 2.1.2. Neurocranium 2.1.2.1. Frontal. The frontal is in union with its mate along its length by way of a strong, interdigitate suture. In lateral view, the frontals form a dome, the apex of which is posterior to the midpoint of that element (Fig. 2). Frontal doming is typical of juvenile lambeosaurines, but is present also in Lophorhothon (Horner et al., 2004) and juvenile Saurolophus angustirostris (Bell, 2011). The anterior margin is slightly upturned (but is lower than the apex of the frontal dome) where it contacts the nasal. Each frontal has a wide embayment, which receives the nasal. Consequently, the naso-frontal suture is smoothly W-shaped in posterodorsal view (Fig. 2), similar to Jaxartosaurus (PIN 1/5009) and juvenile Hypacrosaurus (Evans, 2010, fig. 2-3), but contrasting with the square to U-shaped suture in juvenile Corythosaurus, Lambeosaurus, H. stebingeri, and Velafrons (Evans et al., 2005, 2007; Brink et al., 2011). The sutural surface is anteroventrally oriented in lateral view; however, there is no evidence of a frontal promontorium as is present in Parasaurolophus and Charonosaurus (Godefroit et al., 2001; Evans et al., 2007). The lateral margin of the frontal abuts the prefrontal anteriorly and the postorbital posteriorly. The frontaleprefrontal suture is oriented anteromedially and the frontalepostorbital suture is posteromedially oriented in dorsal aspect. This union excludes the frontal from the orbital rim. Ventrally, the frontal is contacted from anterior to posterior by the presphenoid, orbitosphenoid, and laterosphenoid. Together, these elements enclose the cerebral hemispheres and olfactory tract. Lateral to the frontalepresphenoideorbitosphenoid suture, the frontal is concave forming the dorsomedial wall of the orbital cavity. 2.1.2.2. Parietal. The paired parietals form a single medial element that ossifies early in ontogeny to roof the cerebellar cavity. They are wider than long and saddle-shaped in lateral view. In dorsal aspect, the lateral walls of the parietals are concave. The anterior sutural surface with the frontals is broadly M-shaped in anterodorsal view; a broad triangular process of the parietals separates the frontals for a short distance anteriorly (Fig. 2). Ventrally, the parietals form a straight suture with the prootic and with the laterosphenoid for a short distance more anteriorly. Posteriorly, the squamosals exclude the parietals from the posterior margin of the skull. This feature is absent in Jaxartosaurus (PIN 1/5009) and questionably absent in Aralosaurus (PIN 2229), but can be variable in lambeosaurines (Brink, 2009). A narrow sagittal ridge on the posterodorsal half of the parietals is continuous with the sagittal crest formed by the squamosals more posteriorly. 2.1.2.3. Presphenoid. The paired presphenoids form the anteroventral part of the neurocranium (Fig. 3). The presphenoid is concave laterally where it forms the dorsomedial wall of the orbital cavity. It forms sutures with the frontal dorsally, the orbitosphenoid posteriorly, and its compliment ventromedially. Together, the 5 presphenoids are Y-shaped in anterior view; the dorsal V-shaped half transmits the neural olfactory system. Ventral to this point, the presphenoids meet to form the interorbital septum; the presphenoidepresphenoid suture is clearly visible in anterior view. The presphenoid appears to be broken ventrally and does not contact the basisphenoid. 2.1.2.4. Orbitosphenoid. The dorsoventrally elongate orbitosphenoid contacts the frontal dorsally, the laterosphenoid posteriorly, and the basisphenoid ventrally. The orbitosphenoid is divided into dorsal and ventral components divided by a deep, horizontal invagination of the anterior border of the element (Fig. 3). This invagination is unlike any other hadrosaurid orbitosphenoid, but this region is not particularly well preserved in Kazaklambia and might in fact be broken. The orbitosphenoid contacts the presphenoid dorsal to this invagination whereas it is free below this point. The optic nerve (cranial nerve [c.n.] II) likely exited from a foramen at the posterior end of the invagination; however, there is no evidence of the foramen for the trochlear nerve (c.n. IV), which typically pierces the orbitosphenoid in other hadrosaurids (Evans, 2010). With the laterosphenoid and basisphenoid, the posteroventral corner of the orbitosphenoid forms the anterodorsal margin of the foramen for the abducens nerve (c.n. VI). 2.1.2.5. Laterosphenoid. In lateral view, the laterosphenoid is vertically oriented and rises above the level of the prootic. The dorsal head of the laterosphenoid inserts into a cotylus on the ventral surface of the postorbital forming a joint that was likely synovial in nature. The crista antotica is nearly confluent with the anterior border of the laterosphenoid, extending dorsoventrally along the dorsal half of the laterosphenoid. Ventrally, this ridge turns posteroventrally, terminating approximately halfway between the openings for the trigeminal (c.n. V) and abducens (c.n. VI) nerves. This same ridge probably delimits the dorsal margin of passage for the ophthalmic branch of the trigeminal nerve (c.n. V). Ventral to this ridge, the alar process is subtriangular in lateral view and closely appressed to the surface of the laterosphenoid (Fig. 3). Immediately ventral to the alar process, the laterosphenoid forms a suture with the basisphenoid. The anterior margin of the laterosphenoid is straight along its contact with the orbitosphenoid. The ventral limit of this contact is interrupted by the opening for the abducens nerve (c.n. VI). The posterior union between the laterosphenoid and the prootic is largely obscured by fusion. 2.1.2.6. Prootic. Although the margins of the prootic are largely obscured by fusion, its constituent foramina are visible (Fig. 3). Posteriorly, the prootic defines the anterior margin of the common foramen for glossopharyngeal nerve (c.n. IX) and the oval window. The facial nerve (c.n. VII) exits via a small foramen just anterior the opening for c.n. IX. Immediately anterior to the foramen for c.n. VII, a wide embayment of the anterior margin of the prootic defines the posterior half of the large trigeminal foramen. Dorsal and ventral to this foramen, the prootic is indistinguishably fused with the posterior margin of the laterosphenoid. The crista prootica is a horizontal ridge immediately ventral and subparallel to the prooticeparietal suture. 2.1.2.7. Otoccipital. The exoccipital and opisthotic fuse early in embryonic development to form the otoccipital. Together, they form the dorsal and lateral limits of the occiput. The basioccipital processes, which enclose the subcircular foramen magnum, are mediolaterally expanded ventrally where they abut the basioccipital (Fig. 2D). This suture is dorsomedially inclined. The lateral wall of the basisphenoid process is pierced by two foramina, which form a horizontal line posterior to the crista tuberalis. These foramina correspond to the vagus and accessory nerves (c.n. Please cite this article in press as: Bell, P.R., Brink, K.S., Kazaklambia convincens comb. nov., a primitive juvenile lambeosaurine from the Santonian of Kazakhstan, Cretaceous Research (2013), http://dx.doi.org/10.1016/j.cretres.2013.05.003 6 P.R. Bell, K.S. Brink / Cretaceous Research xxx (2013) 1e10 Fig. 3. Braincase of PIN 2230/1 with jugal and postorbital processes removed (cross-hatching). Cranial openings (dark grey) denoted by roman numerals (iev). Matrix is light grey. Approximate field of view equals 11 cm. Abbreviations: Bs, basisphenoid; Cp, cornoid process; Fo, fenestra ovalis; Fr, frontal; Ju, jugal; L, lacrimal; Ls, laterosphenoid; Mpa, medial palatine artery; Or, orbitosphenoid; Pl, palatine; Po, postorbital; Pr, prootic; Pfr, prefrontal; Ps, presphenoid; Q, quadrate; Qf, pterygoid flange of quadrate; Sq, squamosal. X and XI) anteriorly and the hypoglossal nerve (c.n. XII) posteriorly. The crista tuberalis forms the otoccipitalebasioccipital juncture posterodorsally where it is confluent with the ventral margin of the paroccipital process. Anterodorsal to the foramen for c.n. X and XI is a dorsoventrally elongate, figure-eight shaped foramen. The dorsal component forms the oval window and the ventral portion transmits the glossopharyngeal nerve (c.n. IX). The posterior margin of this fenestra is formed by the otoccipital and defines the anterior extent of that bone. Dorsal to this fenestra, the otoccipital is indistinguishably fused with the prootic. Posteriorly, dorsal to the foramen magnum, the otoccipitals meet along a straight, vertical suture that is nearly closed in PIN 2230/1. From this suture, the otoccipital diverts laterally to form the paroccipital process. A prominent horizontal ridge extends from the midline along the posterior face of the paroccipital process at about its midheight. It does not reach the lateral edge of this element. The distal (lateral) part of the paroccipital process is bent ventrolaterally and tapers to a blunt, squared-off terminus. This ventral terminus is approximately level with the ventral margin of the foramen magnum. 2.1.2.8. Basioccipital. Only the occipital condyle of the basioccipital is preserved (Fig. 2D). In posterior view, it is roughly trapezoidal and the ventral margin convex down. To what extent the basioccipital is involved in the formation of the foramen magnum cannot be determined from its incomplete preservation. 2.1.2.9. Basisphenoid. The basisphenoid is observable only through the orbits of PIN 2230/1. Along its dorsal margin, the basisphenoid contacts the orbitosphenoid anteriorly and laterosphenoid posteriorly. The pterygoid flange of the quadrate obscures the relationship of the basisphenoid to the prootic, basioccipital, and otoccipital. Where the basisphenoid, orbitosphenoid, and laterosphenoid converge, they form the foramen for the abducens nerve (c.n. VI). The finger-like cultriform (parasphenoid) process is straight and nearly perpendicular to the (antero) ventral margin of the basisphenoid in lateral view. The cultriform process extends anteriorly to contact the distal ends of the palatine process of the pterygoid. 2.1.3. Palate Elements of the palate are visible only through the temporal fenestrae and in posterior aspect of the skull. As visible, the palatine is triangular and nearly two-thirds the height of the orbit. The posterior edge is nearly vertical. The palatine process of the pterygoid rises anterodorsally at a 45 angle to overlie the posterior edge of the palatine. The palatine process is thin and strap-like and the dorsal edge is enrolled medially. The pterygoids do not contact one another but fuse distally to the palatine. The right pterygoid terminates immediately dorsal to the left pterygoid and is abutted from behind by the anterior tip of the cultriform process. The dorsal and ventral quadrate processes of the pterygoid are visible in posterior view where they form an extensive lap joint with the quadrate (Fig. 2D). The ventral quadrate process is buttressed medially for its entire length. Its distal (posterior) terminus is spatulate and extends from the posterior margin of the element. The dorsal quadrate process is triangular and buttressed along its dorsal edge. Together, the quadrate processes of the pterygoid occupy the middle third of the height of the quadrate. Please cite this article in press as: Bell, P.R., Brink, K.S., Kazaklambia convincens comb. nov., a primitive juvenile lambeosaurine from the Santonian of Kazakhstan, Cretaceous Research (2013), http://dx.doi.org/10.1016/j.cretres.2013.05.003 P.R. Bell, K.S. Brink / Cretaceous Research xxx (2013) 1e10 2.1.4. Mandible The mandible is broken anteriorly, missing most of the dentary anterior to the coronoid process. The coronoid process is at right angles to the alveolar margin, reminiscent of the plesiomorphic condition of Hadrosauroidea (Prieto-Marquez, 2010a). However, this angle might be influenced in part by the incompleteness of the specimen. The post-dentary bones are tightly articulated although those on the right side have slipped out of articulation with the quadrate. The largest element in this association, the surangular, forms the glenoid articulation with the quadrate. The angular is a mediolaterally-flattened strap adhering to the medial surface of the dentary and surangular. Immediately dorsal to the posterior end of the angular, the small mediolaterally-flattened articular contacts the posteromedial end of the surangular. The teeth are poorly preserved; only the posterior-most teeth retain the characteristic rhomboidal lingual surface and median carinae seen in other hadrosaurids. Marginal denticles could not be observed on any teeth. 3. Morphometric analysis Linear morphometrics have been employed in previous analyses of North American hadrosaurines and lambeosaurines to test for taxonomic and allometric differences in growth among contemporaneous species (Dodson, 1975; Evans, 2010; Campione and Evans, 2011). However, such rigorous analyses as these have not been attempted for Eurasian taxa, in part due to small sample size. The most distinguishing and taxonomically significant morphology of lambeosaurine dinosaurs is the cranial crest. Crest development varies ontogenetically, and is genus-specific, even at juvenile growth stages (Dodson, 1975; Evans, 2010). The proportions of the skull roof are also taxonomically significant, probably in response to the development of the cranial crest (when present). A ratio of the length over width of the frontal (length of the external interfrontal suture posterior to the base of the crest over the maximum transverse width of the frontal) has been used in previous analyses to distinguish between hadrosaurids, basal lambeosaurines, and derived lambeosaurines (Godefroit et al., 2004a; Evans and Reisz, 2007; Gates et al., 2007; Godefroit et al., 2008). Although the parameters of the frontal change through ontogeny, differences between the allometric trajectories of distinct taxa are significantly different (Evans et al., 2007; Evans, 2010). Hadrosaurines, Jaxartosaurus, and Amurosaurus all have relatively long frontals, which grow isometrically relative to skull length through ontogeny (Godefroit et al., 2004a). The frontals of lambeosaurines are negatively allometric, remaining proportionately short as the skull grows in Corythosaurus, Hypacrosaurus, and Lambeosaurus (Evans, 2010), while Parasaurolophus and Charonosaurus have relatively the shortest frontals of all known lambeosaurines (Evans et al., 2009; Evans and Reisz, 2007; Evans et al., 2007; Godefroit et al., 2001). To examine the position of Kazaklambia relative to other hadrosaurids, jugal length, development of the crest, and proportions of the frontal were examined with a regression analysis. The dataset used in this analysis (Appendix A) is a compilation of cranial measurements from Evans et al. (2007), Godefroit et al. (2004a,b), and newly collected data, which includes 61 specimens from 12 species of lambeosaurines, five species of hadrosaurines, and one hadrosauroid, from juvenile, subadult, and adult growth stages. Jugal length and crest height above the teeth were regressed against quadrate height as a proxy for skull size for lambeosaurines only, and the residuals were plotted to examine the difference between the actual values and the predicted values for that variable. The frontal length for Aralosaurus and the width and length for Amurosaurus and Bactrosaurus were 7 measured from photographs (Godefroit et al., 1998, 2004a; 2004b) using Image J software (Rasband, 2008). All measurements were log transformed before each analysis to standardize the data. The analysis was completed using the program JMP (2010, Version 9). 3.1. Results The plot of the residuals of the regression of jugal length vs. quadrate height (Log Jugal Length ¼ 0.1582039 þ 0.9075069*Log Quadrate Height, R2 ¼ 0.953) shows that the length of the jugal of Kazaklambia is near what is expected for a juvenile lambeosaurine (Fig. 4A). The values of the residuals for Kazaklambia are similar to subadult skulls of Hypacrosaurus stebingeri (Fig. 4A). The plot of the residuals of the regression of skull height above teeth vs. quadrate height (Log Ht Above Teeth ¼ 1.861624 þ 1.8525295*Log Quadrate Height, R2 ¼ 0.845) shows that the height of the crest in Kazaklambia is higher than what is expected for a juvenile lambeosaurine with a similar quadrate height, such as Hypacrosaurus altispinus, H. stebingeri, Corythosaurus, and Lambeosaurus (Fig. 4B). The results of the regression of frontal length vs. frontal width show a distinction between the frontals of lambeosaurines and hadrosaurines (Fig. 4C). Parasaurolophus has the shortest frontal through ontogeny, with one juvenile specimen falling outside the 95% confidence interval for lambeosaurines, while the frontals of Corythosaurus, Hypacrosaurus, and Lambeosaurus are slightly longer throughout their trajectory. Of the hadrosaurines, Shantungosaurus has the widest frontals, while the frontals of Edmontosaurus, Gryposaurus, and Prosaurolophus are narrower and have similar trajectories through ontogeny. Some frontals of adult specimens of Edmontosaurus and Prosaurolophus converge on lambeosaurine frontal proportions, but remain within the 95% confidence interval for hadrosaurines. Interestingly, Jaxartosaurus plots just outside the 95% confidence interval for lambeosaurines, but within the range of variation for the hadrosaurines. The frontal of Kazaklambia falls just outside the range of variation of hadrosaurines, and is most similar to the frontals of the juvenile Amurosaurus and Tsintaosaurus. 4. Discussion 4.1. Comparison of Aralosaurus tuberiferus and Jaxartosaurus aralensis to Kazaklambia convincens We agree with previous assertions that Kazaklambia is a member of Lambeosaurinae based on the possession of several cranial characters: 1. Hollow narial crest formed by the nasal and premaxilla; 2. Oval supratemporal fenestra with long-axis oriented anterolaterally; 3. Parietal shortened (length/width <2); 4. Proportions of the frontal (length over width <0.8); 5. steeplyangled parietal bending below level of postorbital-squamosal bar; and 6. parietal crest occupying more than half the length of that element (Evans and Reisz, 2007; Prieto-Marquez, 2010a; Gates et al., 2011). Given the clear lambeosaurine affinities of K. convincens, it is not necessary to compare it to basal hadrosauroids, Batyrosaurus rozhdestvenskyi and the holotype of ‘Arstanosaurus akkurganensis’; however, two other lambeosaurines are known from Kazakhstan: Aralosaurus tuberiferus and Jaxartosaurus aralensis. Aralosaurus tuberiferus (PIN 2229) is known from a single partial skull from the Beleutinskaya Svita from the SakheSakh locality in central Kazakhstan. The age of this formation is tentatively regarded as Turonian (Rozhdestvensky, 1974; Godefroit et al., 2004b). Jaxartosaurus aralensis (PIN 1/ 5009) was erected on the basis of the posterior part of one skull, dentary, surangular, and several postcranial elements recovered Please cite this article in press as: Bell, P.R., Brink, K.S., Kazaklambia convincens comb. nov., a primitive juvenile lambeosaurine from the Santonian of Kazakhstan, Cretaceous Research (2013), http://dx.doi.org/10.1016/j.cretres.2013.05.003 8 P.R. Bell, K.S. Brink / Cretaceous Research xxx (2013) 1e10 from the KyrkeKuduk locality (Dabrazinskaya Svita, lower Santonian) in the Chuley region of Chimkent/Tashkent. Only the skull is currently available for study, the remaining elements having been lost (Godefroit et al., 2004b). Both taxa have been described in detail (Rozhdestvensky, 1968; Godefroit et al., 2004b) and it is not necessary to redescribe them here; however, it is necessary to make some comments regarding their relationships with Kazaklambia. Aralosaurus is regarded as the most primitive lambeosaurine and one of the oldest hadrosaurids (Godefroit et al., 2004b; PrietoMarquez, 2010a,b; but see Sues and Averianov, 2009 for another interpretation). Rozhdestvensky (1968) originally regarded Aralosaurus as a gryposaur-like hadrosaurine, a sentiment that was upheld until recently. Godefroit et al. (2004b) rediagnosed Aralosaurus as a basal lambeosaurine citing a suite of primitive characters that differentiate it from other lambeosaurines, including Kazaklambia: 1) nasal terminates anterior to the orbit; 2) transverse frontal-nasal contact in dorsal view; 3) frontals participate in orbital margin and; 4) fronto-nasal fontanella present (also present in Aralosaurus and Amurosaurus). We agree with Godefroit et al. (2004a) that Jaxartosaurus represents a valid taxon on account of the autapomorphies listed by those authors: 1) lateral bar of the supratemporal fenestra short and robust; and 2) alar process (prootic process) of the laterosphenoid thickened. These characters suggest that Kazaklambia is not a juvenile of Jaxartosaurus, as previously suspected (Weishampel and Horner, 1990; Horner et al., 2004). In addition, Jaxartosaurus comes from a different formation and lower stratigraphic level (Santonian) than Kazaklambia, although well-constrained ages for both levels are unknown. Perhaps the most salient feature is the medial ramus of the squamosal, which is horizontal in Jaxartosaurus and unlike the dorsomedially oriented rami in other lambeosaurines (Godefroit et al., 2004a, b; Evans, 2010), including Kazaklambia. The robust alar process of the laterosphenoid seen in Jaxartosaurus contrasts with the flattened and more gracile process in Kazaklambia. In addition, the ophthalmic branch of the trigeminal nerve in Jaxartosaurus was borne along a deep anteroposteriorly-directed sulcus that bisects the lateral face of the laterosphenoid. In Kazaklambia, the path of the ophthalmic branch is delimited only by single prominent ridge along the anterior part of the laterosphenoid and there is no sulcus. Anteriorly, the presphenoid of Jaxartosaurus is anteroposteriorly short, forming a strap-like element in lateral view that is roughly one quarter the length of the orbitosphenoid. The presphenoid of Kazaklambia, although imperfectly preserved, is at least as long as the orbitosphenoid and consequently roughly equidimensional. The prefrontal dome is present over the orbits in Kazaklambia, lateral to the frontal dome (Fig. 2). By comparison, the frontal dome is weakly expressed in Jaxartosaurus (although this is almost certainly growth related), whereas the postorbital dome is entirely absent. An Asian origin for Lambeosaurinae has been convincingly argued in recent years (Godefroit et al., 2004b; Prieto-Marquez, 2010b). Conveniently, the most primitive lambeosaurines known to date (the Asian taxa Aralosaurus, Jaxartosaurus, and Tsintaosaurus) are also the oldest. The basal position of Kazaklambia based on the morphometric analysis and its stratigraphic position is consistent also with the stratigraphic and geographic locations of other primitive lambeosaurines and bolsters support for an Asian origin of this clade. Fig. 4. Morphometric analysis of hadrosauroid dinosaurs. A, residual plot for regression of jugal length against quadrate height for lambeosaurines. B, residual plot for regression of skull height above teeth against quadrate height for lambeosaurines. C, regression of frontal length against frontal width for hadrosauroids. The 95% confidence ellipses are for Lambeosaurinae (left) and Hadrosaurinae plus Bactrosaurus (right). 4.2. Juvenile status and affinities of Kazaklambia convicens As the only known specimen of Kazaklambia is a juvenile, it was not appropriate to include it in a phylogenetic analysis of Please cite this article in press as: Bell, P.R., Brink, K.S., Kazaklambia convincens comb. nov., a primitive juvenile lambeosaurine from the Santonian of Kazakhstan, Cretaceous Research (2013), http://dx.doi.org/10.1016/j.cretres.2013.05.003 P.R. Bell, K.S. Brink / Cretaceous Research xxx (2013) 1e10 Hadrosauridae, as juvenile taxa are infrequently recovered in a clade with their corresponding adult morph (Fowler et al., 2011; Tsuihiji et al., 2011; Campione et al., 2013). The ability to compare species at different ontogenetic stages, especially with the development of elaborate cranial crests, is extremely difficult. Therefore, the diagnosis for this taxon is based on characters that are only present at this ontogenetic stage, and suggest that it is different from other lambeosaurine taxa based on a differing growth trajectory. The morphometric comparison of skull size and crest height suggests that for its size, Kazaklambia has a better-developed crest than that of North American lambeosaurines (Fig. 4A and B). This could potentially indicate a similarity to parasaurolophs, where the crest is suggested to develop early in ontogeny (Evans et al., 2007, 2009). The doming of the postorbitals could also be indicative of a relationship between parasaurolophs and Kazaklambia, as the postorbitals of the parasauroloph Charonosaurus are also domed at immature growth stages, and probably change with the development of the enlarged, tubular cranial crest (Godefroit et al., 2001). Recent work on North American lambeosaurines has shown the morphology of the skull roof is diagnostic even at an early ontogenetic stage. The frontal of Parasaurolophus presents a steeply-angled frontal-nasal sutural surface, which appears peculiar to this taxon (Evans et al., 2007). The frontals of juvenile corythosaurins (Corythosaurus, Hypacrosaurus, and Lambeosaurus) are virtually identical; however, Hypacrosaurus can be differentiated on account of its relatively short parietal, a feature also shared with Parasaurolophus (Evans, 2010). Morphometric analysis in this study demonstrates the frontals of Kazaklambia are more elongate than those of other lambeosaurines, both adults and juveniles, and especially parasaurolophs. Therefore, the relative proportions of the skull roof should be sufficient to identify adult specimens of Kazaklambia when they are found. The proportions of the frontal of Kazaklambia are also similar to the frontals of Amurosaurus and Tsintaosaurus, even though the holotype of Tsintaosaurus is considered to be adult (Buffetaut and TongBuffetaut, 1993). This suggests that at the juvenile growth stage, Kazaklambia falls within the range of variation of the frontal for basal lambeosaurines, whereas it is noticeably different from the juveniles of Parasaurolophus sp., Hypacrosaurus spp., Corythosaurus spp., and Lambeosaurus spp. from North America. With more specimens, the ontogenetic trajectories of the Eurasian taxa may be compared to each other and to the North American taxa, and the similarities between Kazaklambia and parasaurolophs can be further explored. 5. Conclusions North American species of the genus Procheneosaurus are unanimously regarded as junior synonyms of Corythosaurus spp., Lambeosaurus spp., and Hypacrosaurus stebingeri (Dodson,1975; Weishampel and Horner, 1990; Horner and Currie, 1994; Horner et al., 2004; Evans et al., 2005). We therefore present a new generic moniker for ‘Procheneosaurus’ convincens, Kazaklambia convicens comb. nov., from the Santonian of Kazakhstan. Kazaklambia is one of the earliest known lambeosaurines, and along with Aralosaurus and Jaxartosaurus, also from Kazakhstan, supports an Asian origin for Lambeosaurinae (Godefroit et al., 2004a,b, 2008; Prieto-Marquez, 2010b). A suite of anatomical features differentiate K. convincens from both Aralosaurus and Jaxartosaurus and further suggest that adult specimens should be identifiabledparticularly with reference to the elongate frontaldwhen they are found. Although the precise phylogenetic position of Kazaklambia is unknown, morphologic (particularly the crest-forming elements and proportions of the frontal) evidence argues for a close alliance with basal lambeosaurines. 9 Acknowledgements We thank V. Alifanov and T. Tumanova (PIN, Moscow) for their hospitality and access to specimens in their care, D. Evans (ROM) for discussion, and to N. Campione (University of Toronto) for access to unpublished data. Terry Gates and three anonymous reviewer provided useful comments on the manuscript. A Dinosaur Research Institute travel grant to PRB and NSERC to KSB are gratefully acknowledged. References Averianov, A.O., Nessov, L., 1995. A new Cretaceous mammal from the Campanian of Kazakhstan. Neues Jahrbuch für Geologie und Paläontologie, Montschefe 1995, 65e74. Bell, P.R., 2011. Cranial osteology and ontogeny of Saurolophus angustirostris from the Late Cretaceous of Mongolia with comments on Saurolophus osborni from Canada. 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Proceedings of the Royal Society B: Biological Sciences 276, 2549e2555. Tsuihiji, T., Watabe, M., Tsogtbaatar, K., Tsubamoto, T., Barsbold, R., Suzuki, S., Lee, A.H., Ridgely, R., Kawahara, Y., Witmer, L.M., 2011. Cranial osteology of juvenile specimens of Tarbosaurus bataar (Theropoda, Tyrannosauridae) from the Nemegt Formation (Upper Cretaceous of Bugin Tzav, Mongolia). Journal of Vertebrate Paleontology 31, 497e517. Weishampel, D.B., Horner, J.R., 1990. Hadrosauridae. In: Weishampel, D., Dodson, P., Osmólska, H. (Eds.), The Dinosauria, second ed. University of California Press, Berkeley, pp. 534e561. Appendix A. Supplementary data Supplementary data related to this article can be found at http://dx.doi.org/10. 1016/j.cretres.2013.05.003. Please cite this article in press as: Bell, P.R., Brink, K.S., Kazaklambia convincens comb. nov., a primitive juvenile lambeosaurine from the Santonian of Kazakhstan, Cretaceous Research (2013), http://dx.doi.org/10.1016/j.cretres.2013.05.003