Osteology of Zalmoxes shqiperorum (Dinosauria, Ornithopoda), based on new specimens from the Upper Cretaceous of Na˘ lat,-Vad (Romania)

Pascal GODEFROIT Département de Paléontologie, Institut royal des Sciences naturelles de Belgique, 29 rue Vautier, B-1000 Bruxelles (Belgium) [email protected]

Vlad CODREA Catedra de Geologie, Facultatea de Biologie şi Geologie, Universitatea din Babes-Bolyai, 1 Kogalniceanu st., RO-3400 Cluj-Napoca (Romania) [email protected]

David B. WEISHAMPEL Center for Functional Anatomy & Evolution, School of Medicine, Johns Hopkins University, 1830 E Monument st. RM 303, Baltimore MD 21205 (USA) [email protected]

Godefroit P., Codrea V. & Weishampel D. B. 2009. — Osteology of Zalmoxes shqiperorum (Dinosauria, Ornithopoda), based on new specimens from the Upper Cretaceous of Na˘ lat,- Vad (Romania). Geodiversitas 31 (3) : 525-553.

ABSTRACT Nălaţ-Vad is a new fossil locality discovered in 2002 in the Sănpetru Formation (Maastrichtian, Late Cretaceous) of the Haţeg Basin (Transylvania, Romania). Th is site has, among others, yielded the most complete skeleton that can be referred to the ornithopods dinosaur Zalmoxes shqiperorum Weishampel, Jianu, Cziki & Norman, 2003, but also isolated elements belonging to both juveniles and adult individuals. Th is material provides new information about the anatomy KEY WORDS Dinosauria, of Z. shqiperorum, and about the inter- and intraspecifi c variability within Zal- Ornithopoda, moxes. Zalmoxes robustus (Nopsca, 1902) and Z. shqiperorum were apparently Zalmoxes shqiperorum, sympatric species in Transylvania by latest Cretaceous time. Th e co-existence Upper Cretaceous, Haţeg Basin, in the same locality of two closely-related species is not an isolated case among sympatry. ornithopod dinosaurs.

GEODIVERSITAS • 2009 • 31 (3) © Publications Scientifi ques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 525 Godefroit P. et al.

RÉSUMÉ Ostéologie de Zalmoxes shqiperorum (Dinosauria, Ornithopoda), d’après de nou- veaux spécimens du Crétacé supérieur de Nălaţ-Vad (Roumanie). Nălaţ-Vad est un nouveau site fossilifère découvert en 2002 dans la Formation de Sănpetru (Maastrichtien, Crétacé supérieur) du bassin de Haţeg (Transylvanie, Roumanie). Ce site a notamment livré le squelette le plus complet à ce jour pou- vant être attribué au dinosaure ornithopode Zalmoxes shqiperorum Weishampel, Jianu, Cziki & Norman, 2003, mais également des ossements isolés appartenant à des individus juvéniles et adultes. Ce matériel apporte de nouvelles informations MOTS CLÉS Dinosauria, sur l’anatomie de Z. shqiperorum, mais également sur la variabilité inter- et intra- Ornithopoda, spécifi que au sein même du genre Zalmoxes. Zalmoxes robustus (Nopsca, 1902) et Zalmoxes shqiperorum, Z. shqiperorum étaient apparemment des espèces sympatriques en Transylvanie Crétacé supérieur, bassin de Haţeg, au Crétacé supérieur. La coexistence dans une même localité de deux espèces sympatrie. proches n’est pas un cas isolé chez les dinosaures ornithopodes.

INTRODUCTION Mochlodon robustum Nopcsa, 1902 (Nopcsa 1902, 1904). He later suggested that Mochlodon might be Although Rhabdodontidae probably comprises identical to Rhabdodon and that sexual dimorphism the most abundant dinosaurs from the Upper might be responsible for the observed diff erences Cretaceous of Europe, these medium-sized ornitho- between the two forms (Nopcsa 1915, 1929). pods remained poorly understood until recently. Ösi (2004) reported rhabdodontid fossils in the Rhabdodon priscus Matheron, 1869 was described Upper Cretaceous of Hungary. Weishampel et al. from fragmentary material discovered in the lower (2003) recently reviewed the rhabdodontid mate- Maastrichtian of La Nerthe (Bouches-du-Rhône, rial from Transylvania, and referred it to as a new France). Additional material from diff erent locali- genus, Zalmoxes. Th ey distinguished two species: ties in southern France, referred to as R. priscus, Z. robustus (Nopcsa, 1902), known from abun- was described later by Lapparent (1947), Garcia dant skull and postcranial material, and the more et al. (1999), Pincemaille-Quillévéré (2002), and poorly known species Z. shqiperorum Weishampel, Pincemaille-Quillévéré et al. (2006). Buff etaut & Jianu, Cziki & Norman, 2003. Both species are Le Loeuff (1991) described Rhabdodon septimani- known from several localities in the Haţeg Basin, cus from a dentary discovered in the Campano- but also from the Vintu de Jos area (Fig. 1). Th e Maastrichtian of Montmouliers (Hérault, France). holotype of Z. shqiperorum (BMNH R4900) is a Rhabdodon Matheron, 1869 specimens have also single incomplete adult individual. Much of the been described in diff erent Campano-Maastrichtian referred material pertains to a partially associated localities from northern Spain (Pereda-Suberbiola & skeleton of a juvenile, which includes most of the Sanz 1999). Bunzel (1871) described Iguanodon pelvis and associated dorsal, sacral and caudal ver- suessi from the Upper Cretaceous of Niederöster- tebrae (FGGUB R1087-1133 and R1355-1357). reich (Austria). Seeley (1881) subsequently created From the skull of Z. shqiperorum, only the dentary the genus Mochlodon to include this species. In the is known to date. Both the fore- and hindlimbs beginning of the 20th century, Nopcsa described were also incompletely known. Sachs & Hornung important ornithopod material from the Upper (2006) referred specimens discovered in Austria Cretaceous of the Haţeg Basin in Transylvania. Part to Zalmoxes. of this material was referred to Mochlodon suessi New material that can be referred to as Z. shqipero- Seeley, 1881 and another part to a new species rum was discovered in 2002 at Nălaţ-Vad, a new

526 GEODIVERSITAS • 2009 • 31 (3) Osteology of Zalmoxes shqiperorum (Dinosauria, Ornithopoda) from Romania

locality in the Sănpetru Formation of the Haţeg Metaliferi mountains Mures Basin. Th e new material, housed in the Catedra de Cugir Geologie, Facultatea de Biologie şi Geologie, Uni- Orastie versitatea din Babes-Bolyai (Cluj-Napoca, Roma- Poiana Rusca mountains nia), includes the most complete skeleton that can Orastie hills II e be referred to date to this taxon (UBB NVZ1), but Govajdia

rasti also isolated elements belonging to both juvenile and O

Sureanu Strei

Hunedoara hills adult individuals. It provides new information about mountains the anatomy of Z. shqiperorum, and about the inter- Cerna Valioara Tustea Hateg‚ and intraspecifi c variability within Zalmoxes. Rusca Densus Nalat˘ ‚ Vad Th e aim of the present paper is to complete the osteo- I Montana˘ Totesti e logical description of Z. shqiperorum, whose skeleton Sanpetru˘ Pui Mar is scarcely known so far, on the basis of the newly- III Râul

l discovered material from Nălaţ-Vad. Th e diagnoses of Petrosani Cluj-Napoca Sibise ‚ Barbat both Zalmoxes and Z. shqiperorum and the apparent IV sympatry of two closely-related species of medium- Hateg‚ Retezat Bucharest mountains Jiul de Vest sized herbivorous dinosaurs will be discussed. 15 km

ABBREVIATIONS FIG. 1. — Geographical location of the different localities that have BMNH Department of Palaeontology, Natural His- yielded fossils of Zalmoxes robustus (Nopsca, 1902) (black stars) tory Museum, London; and Z. shqiperorum Weishampel, Jianu, Cziki & Norman, 2003 (white FGGUB Facultatea de Geologie şi Geofi zics, Univer- stars) in Haţeg and surrounding basins (Romania): I, Rusca Montană sitatea din Bucareşti; Basin; II, Strei Basin; III, Haţeg Basin; IV, Petroşani Basin. UBB Catedra de Geologie, Facultatea de Biologie şi Geologie, Universitatea din Babes-Bolyai, Cluj-Napoca. Numerous palaeosols developed within these de- posits, as indicated by the presence of calcretes and GEOLOGICAL SETTING the reddish coloration of the sediments (Smith et al. 2002; Van Itterbeeck et al. 2004). Dinosaur eggs were Th e Nălaţ-Vad locality is a newly discovered exposure found in these calcretes, but they are less numerous situated in the bedding of the Raul Mare River between than at Toteşti-Baraj. On the other hand, vertebrate the villages of Nălaţ and Vad. Usually, the water fl ow remains are by far more numerous at Nălaţ-Vad. Al- covers nearly the entire exposure, so that the excavation though isolated bones were randomly found in the conditions are particularly hard. Th e strike of the layers entire locality, three pockets have yielded concentrated (N40E75N) does not signifi cantly diff ers from the vertebrate remains. A pocket with microvertebrates strike of the Toteşti-Baraj locality (Codrea et al. 2002), consists of a limestone mainly composed of eggshell situated 3 km upstream. According to the geological debris, a so-called coquina that grades into black- map of the area, the outcropping sediments belong coloured marls that is less enriched in egg shells. to the Sănpetru Formation (Grigorescu 1983, 1992; Both the marl and limestones are extremely rich in Grigorescu et al. 1999; Van Itterbeeck et al. 2004). microvertebrate remains, mainly mammals, birds Palaeomagnetic studies suggest that this formation and small theropods. Th e second pocket was rich should entirely belong to the lower Maastrichtian in sauropod bones. Th e third pocket has yielded (Panaiotu & Panaiotu 2002). Th e facies assemblage disarticulated Zalmoxes bones, apparently belonging at Nălaţ-Vad indicates a fl uvial palaeoenvironment to a single individual (UBB NVZ1). However, one with sandy channel infi lls and dominantly black to slender quadrate from this pocket apparently does reddish-brown overbank fi nes. Th e overbank fi nes not belong to Zalmoxes, more closely resembling that are dominantly silts and clays representing suspen- of the more advanced iguanodontian Telmatosaurus sion deposits on the ancient fl oodplain. Nopcsa, 1903.

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SYSTEMATIC PALAEONTOLOGY angular buccal emargination that forms a horizontal platform extending for the full length of the dentition behind and medial to the coronoid process; scapular Order ORNITHISCHIA Seeley, 1887 blade narrow, strap-like proximally, expanding sharply Suborder ORNITHOPODA Marsh, 1881 posterodistally; expanded region of the scapula adja- EUORNITHOPODA cent to the coracoid suture; acromial process forming (sensu Weishampel 1990) a prominent fl ange; deltopectoral crest of the humerus Infraorder IGUANODONTIA Sereno, 1986 particularly prominent, extending over the proximal half of the humerus; iliac peduncle of the ischium parti- Family RHABDODONTIDAE cularly developed; distal end of the ischium forming a Weishampel, Jianu, Cziki & Norman, 2003 boot-like expansion.

Genus Zalmoxes DESCRIPTION OF THE ZALMOXES MATERIAL Weishampel, Jianu, Cziki & Norman, 2003 FROM NA˘ LAT,-VAD Th is description is mainly based on UBB NVZ1, TYPE SPECIES. — Zalmoxes robustus (Nopcsa, 1902) by the disconnected skeleton discovered in the third original designation. fossiliferous pocket at Nălaţ-Vad. Approximately EMENDED DIAGNOSIS. — Unambiguous synapomorphies 70% of the skull is known from disarticulated bones of the genus include the following characters: reduced in this specimen. A tentative reconstruction of the external mandibular fenestra positioned anteriorly along skull of UBB NVZ1 is presented in Figure 2A, B. the upper border of the surangular; well-developed supra- acetabular process on the ilium; absence of an obturator Th e axial skeleton, the pes and the manus are very process on the ischium; arched ischial shaft. incompletely preserved in UBB NVZ1. Th erefore, Th e following characters may be synapomorphic for Z. ro- the description of the postcranium is completed by bustus and Z. shqiperorum, but their presence in Rhabdodon information observed on isolated bones collected remains unknown (see discussion): extensive, complex squa- in other places of the Nălaţ-Vad locality. A recon- mose suture between quadratojugal and jugal; post-temporal struction of the whole skeleton of Z. shqiperorum foramen transmitted through the body of the squamosal; curved shelf on the lateral surface of the postorbital; lateral (with the exception of the pes and the manus) is splaying of the quadrate; deep predentary; long, dorsoven- presented in Figure 2C. trally-narrow, twisted preacetabular process. Th e following characters cannot be observed either in Jugal (Fig. 3) (UBB NVZ1-32) Z. shqiperorum or in the diff erent species of Rhabdodon. Th e jugal of Z. shqiperorum is more gracile than that Th erefore, subsequent discoveries may shift their dis- tribution either inclusively (as synapomorphies for the of Z. robustus (see Weishampel et al. 2003: fi g. 6A, Rhabdodontidae clade) or exclusively (as autapomorphies B). Th e dorsal border of the long maxillary proc- for Z. robustus): absence of the scar for m. adductor man- ess forms a very wide platform-like ventral orbital dibulae externus superfi cialis on the squamosal; large, disc- rim (Fig. 3B). Th e lateral surface of the maxillary shaped quadratojugal; frontal with complex transverse process bears an elongated groove (Fig. 3A). Quite sutural surface that extensively overlaps the parietal; high lateroventral processes on the predentary. anteriorly, the external side of the maxillary process forms a small but deep recess for articulation with the posteroventral corner of the lacrimal. Th e medial Zalmoxes shqiperorum side of the jugal forms a very elongated and deep Weishampel, Jianu, Cziki & Norman, 2003 recess for articulation with the posterior part of the maxilla. At the level of the posteroventral corner of HOLOTYPE. — BMNH R.4900, left dentary, sacrum, the orbit, the medial side of the maxillary process right scapula, right coracoid, partial ?left ilium, partial right ilium, right ischium, left distal ischium and left bears a wide grooved and ridged facet for intimate femur. articulation with the ectopterygoid (Fig. 3C). Th e posterior process is very compressed mediolaterally. EMENDED DIAGNOSIS. — Species of Zalmoxes charac- terised by the following autapomorphies: occipital con- Its ventral border is sigmoid, thickened and cov- dyle not separated from the sphenooccipital tubercles ered with longitudinal striations, indicating fi brous (= basal tubera) by a distinct neck; dentary with an attachment of the superfi cial tissues associated with

528 GEODIVERSITAS • 2009 • 31 (3) Osteology of Zalmoxes shqiperorum (Dinosauria, Ornithopoda) from Romania

B ext.n

A ptf f po na sq pf pop itf pf la na la orb q ju orb ext.n qj

f mx ju pmx po saf itf

pd dt sa an stf sq p

C

FIG. 2. — Zalmoxes shqiperorum Weishampel, Jianu, Cziki & Norman, 2003, skull reconstruction in left lateral (A) and dorsal (B) views; skeletal reconstruction (except the hands and the feet) in right lateral view (C). Abbreviations: an, angular; dt, dentary; ext.n, external naris; f, frontal; itf, infratemporal fenestra; ju, jugal; la, lacrimal; mx, maxilla; na, nasal; orb, orbital cavity; p, parietal; pd, predentary; pf, prefrontal; pmx, premaxilla; po, postorbital; pop, paroccipital process; ptf, post-temporal fenestra; q, quadrate; qj, quadratojugal; sa, surangular; saf, surangular fenestra; sq, squamosal; stf, supratemporal fenestra. Scale bars: A, B, 5 cm; C, 20 cm. the cheek region. On the medial side of the pos- described by Weishampel et al. (2003) in Z. robus- terior process, a prominent vertical ridge limits a tus. As it has not yet been observed in any other wide articular facet for the quadratojugal (Fig. 3C). ornithopod known to date, it can be considered Th e posterior edge of the jugal is ventrally notched as diagnostic for Zalmoxes. However, the jugal has and forms a slot that received the anterior edge of not been described in Rhabdodon yet, thus it is also the quadratojugal. Th is complex articulation be- possible that this type of articulation characterizes tween the jugal and quadratojugal was previously the rhabdodontid clade.

GEODIVERSITAS • 2009 • 31 (3) 529 Godefroit P. et al.

B posterior process posterior process posterior process

orbital rim infratemporal infratemporal fenestra fenestra lacrimal ectopterygoid recess quadra- quadra- orbital rim facet tojugal tojugal facet recess

external groove C A maxillary recess maxillary process

FIG. 3. — Right jugal of Zalmoxes shqiperorum Weishampel, Jianu, Cziki & Norman, 2003 (UBB NVZ1-32), from the Upper Cretaceous of Nălaţ-Vad (Romania), in lateral (A), dorsal (B), and medial (C) views. Scale bar: 5 cm.

Squamosal (Fig. 4A, B) (UBB NVZ1-33) fenestra forms a large notch between the ventral Th e parietal process of the squamosal is proportion- border of the postorbital process and the anterior ally shorter and higher than in Z. robustus. Its dorsal margin of the squamosal body. Th is notch appears margin, which participates in the posterolateral part lower, but deeper than in Z. robustus (Weishampel of the supratemporal fenestra, is particularly thick et al. 2003: fi g. 9). and rugose. Its medial surface bears a long horizontal Ventrally, the squamosal body forms a large and ridge-bound facet, interpreted by Weishampel et al. deep cotylus for the head of the quadrate. As pre- (2003) as an articulation surface for the paroccipital viously noted by Weishampel et al. (2003), the process. A diff erent interpretation is proposed herein. shape and orientation of the cotylus suggests an In UBB NVZ1-33, the paired squamosals and the oblique orientation for the long axis of the quadrate parietal are still articulated (Fig. 4C). It is clear, from (Fig. 4B). An anteroventrally-pointing prequadratic this specimen, that the ridge-bound facet on the process limits the cotylus anteriorly. On the other medial side of the parietal process of the squamosal hand, the postquadratic process is absent and the intimately contacted the posterolateral surface of posteroventral corner of the squamosal curves medi- the parietal instead. Th e parietal processes of the ally instead, as previously described in Z. robustus paired squamosals nearly contact each other along (Weishampel et al. 2003). On the medial surface of the midline on the occipital aspect of the skull, the squamosal body, a vertical ridge starts from the being only separated by the narrow posteroventral dorsal margin of the cotylus. It can be interpreted keel of the parietal (Fig. 4C, D). as the insertion area for the m. adductor mandibulae Th e postorbital process of the squamosal is also externus medialis. more robust than in Z. robustus. Its lateral surface As in Z. robustus, a foramen, interpreted as the forms a wide and scarred articular facet for the post-temporal fenestra that has migrated from a postorbital (Fig. 4A). Because of the preservation marginal position (Weishampel et al. 2003), perfo- state of the postorbital ramus, it cannot be checked rates the body of the squamosal. In Z. shqiperorum, whether a scar for the origin of m. adductor mandi- this foramen is located at mid-distance between the bulae externus superfi cialis is developed on the squa- notch for the infratemporal fenestra and the pos- mosal of Z. shqiperorum. Th e absence of this scar is terior border of the squamosal body (Fig. 4A, B), regarded by Weishampel et al. (2003) as diagnostic whereas it is apparently positioned more posteriorly for Zalmoxes, but the state of this character remains in Z. robustus. As this foramen is now described unknown in Z. shqiperorum and in Rhabdodon as both in Zalmoxes robustus and in Z. shqiperorum, well. Th e posterior margin of the infratemporal and has not been described in any other ornithopod

530 GEODIVERSITAS • 2009 • 31 (3) Osteology of Zalmoxes shqiperorum (Dinosauria, Ornithopoda) from Romania

parietal process A B supratemporal supratemporal fenestra fenestra

postorbital parietal suture suture

post-temporal postorbital fenestra postorbital process process infratemporal infratemporal fenestra fenestra

cotylus of cotylus of quadrate prequadratic quadrate prequadratic process process

C sagittal crest E

parietal

D left squamosal right squamosal

supraoccipital

FIG. 4. — Zalmoxes shqiperorum Weishampel, Jianu, Cziki & Norman, 2003 (UBB NVZ1-33) from the Upper Cretaceous of Nălaţ-Vad (Romania), left squamosal in lateral (A) and medial views (B); photograph (C) and line drawing (D) of the contact area between the squamosals, the parietal, and the supraoccipital; parietal (E) in dorsal view. Scale bars: A-D, 4 cm; E, 2 cm. known to date, it can therefore be regarded as a in fact synapomorphic for Rhabdodontidae (sensu good synapomorphy for Zalmoxes. Nevertheless, Weishampel et al. 2003), because no squamosal is it cannot be excluded that this character would be currently described in Rhabdodon.

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head head squamosal suture A B

quadratojugal suture

jugal wing jugal wing

CDE F

pterygoid pterygoid wing wing

medial medial condyle lateral condyle condyle lateral condyle

FIG. 5. — Zalmoxes shqiperorum Weishampel, Jianu, Cziki & Norman, 2003 from the Upper Cretaceous of Nălaţ-Vad (Romania), dor- sal part of right quadrate (UBB NVZ1-39) in lateral (A) and medial (B) views; dorsal part of left quadrate (UBB NVZ1-40) in lateral (C), posterior (D), medial (E), and anterior (F) views. Scale bar: 5 cm.

Quadrate (Fig. 5) (UBB NVZ1-39 and NVZ1-40) Th e ventral portion of the quadrate is asym- Only the dorsal part of the right quadrate (Fig. 5A, metrically developed into a large lateral condyle, B) and the ventral portion of the left quadrate which articulated with the surangular part of the (Fig. 5C-F) are preserved in the fossiliferous lens. mandibular glenoid, and a smaller medial condyle, Th e quadrate of Z. shqiperorum is very massive which probably articulated with the articular part in comparison with other ornithopods, closely of the mandibular glenoid (Fig. 5D, F). Weisham- resembling that described in Z. robustus (Wei- pel et al. (2003) showed that the important asym- shampel et al. 2003: fi g. 7). Th ere is no indication metry of the ventral condyles, in association with of a vertical buttress that posteriorly overhangs the orientation of the squamosal cotylus, implies a the main shaft, as it is observed in Z. robustus strongly oblique orientation of the quadrate relative (Weishampel et al. 2003: fi g. 7). At this level, the to the sagittal plane of the skull and a triangular posterior margin of the quadrate shaft is smoothly aspect of the skull in posterior view. Th is character concave (Fig. 5A, B). On the other hand, the is regarded as diagnostic for Zalmoxes, because it is anterolateral border of the quadrate forms a very shared by both species Z. robustus and Z. shqipero- massive jugal wing. rum. However, it must be noted that the important

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asymmetry of the distal quadrate condyles also Th e olfactory bulbs form elongated sulci along the characterises the more derived hadrosaurid clade interfrontal suture. Between the impressions for the (Weishampel et al. 1993; Godefroit et al. 1998). cerebrum and the olfactory bulb, divergent ridges Although it is incompletely preserved, the pterygoid mark articulation with the sphenethmoid plate. wing seems massive. Postorbital (Fig. 7) (UBB NVZ1-41) Parietal (Fig. 4E) (UBB NVZ1-33) Th e postorbital of Z. shqiperorum is particularly Only the posterior part of the parietal, which inti- massive. Th e medial border of its anterior plate is mately contacts the medial side of the squamosals, is very thick and rugose for intimate contacts with the preserved in UBB NVZ1-33. As previously observed frontal. Its lateral edge, which participates in the in Z. robustus (Weishampel et al. 2003), the parietal posterodorsal margin of the orbit, is on the other appears particularly narrow and lozenge-shaped in hand rather sharp. Th e surface of the bone is not suf- cross-section. Its dorsal surface bears a high and fi ciently well preserved in order to assess whether it is sharp sagittal crest, much better developed than pierced by foramina, as it is described in Z. robustus in Z. robustus (Weishampel et al. 2003: fi g. 10C). (Weishampel et al. 2003). At the junction with the Th e height of the sagittal crest progressively lessens base of the squamosal process, the dorsal surface of posteriorly, as it slightly bifurcates. the postorbital plate forms a well-developed curved ledge (Fig. 7B), as also observed in Z. robustus and Frontal (Fig. 6) (UBB NVZ1-38) interpreted by Weishampel et al. (2003) as marking In dorsal view, the frontal is subtriangular and it the origin of part of the external mandibular ad- closely resembles that of Z. robustus (Weishampel ductor musculature. Th is ledge was not described et al. 2003: fi g. 8B). Its anterior margin forms a in other ornithopods; it can therefore be regarded broad but shallow articular surface for the nasal as a valid synapomorphy for Zalmoxes. and, maybe, for the prefrontal.Th e posteromedial Th e squamosal process, which participates in the border of the parietal is slightly notched: the anterior anterolateral margin of the supratemporal fenestra end of the parietal therefore probably separated the and in the dorsal margin of the infratemporal fe- paired frontals at this level. Because the posterior nestra, is straight and very compressed mediolater- portion of the frontal is broken off , it cannot be ally. At the junction between the anterior plate, the checked whether the frontal overlapped for some squamosal process and the jugal process, a very extent the parietal. Th is character is regarded by depressed area marks the synovial contact with the Weishampel et al. (2003) as synapomorphic for laterosphenoid (Fig. 7C). Zalmoxes, but the state of this character remains unknown in Z. shqiperorum and in Rhabdodon. Supraoccipital (Fig. 4C) (UBB NVZ1-33) Laterally, the suture with the postorbital is also very Th e posterior surface of the supraoccipital bears long and thick. Th e relationships between the frontal a strong median nuchal crest. Transverse furrows and the prefrontal remain obscure. Weishampel et separate this crest from lateral low bosses, which al. (2003) proposed that the anterior part of the probably form the insertion areas for m. rectus capitis frontal was overlapped by the posterior part of the posterioris. Th e ventral border of the supraoccipital prefrontal, close to the fronto-nasal suture. But it is notched by the dorsal margin of the foramen is also possible that the anterior part of the long magnum. and thick lateral border of the frontal contacted the prefrontal. In any case, it seems that the frontal Basioccipital-basisphenoid (Fig. 8) (UBB NVZ1-40) was completely or nearly completely excluded from In posterior view, the occipital condyle is wide the orbital margin. and reniform. Its articular surface faces quite Th e ventral surface of the frontal is marked by posteriorly. Its concave dorsal side participates in encephalic impressions. Its posterior portion is deeply the fl oor of the foramen magnum. Contrary to excavated by the anterior part of the cerebrum. Z. robustus, the occipital condyle is not supported

GEODIVERSITAS • 2009 • 31 (3) 533 Godefroit P. et al.

?prefrontal olfactory suture bulb nasal suture A B

sphenethmoid postorbital interfrontal suture suture suture

cerebrum

FIG. 6. — Zalmoxes shqiperorum Weishampel, Jianu, Cziki & Norman, 2003 from the Upper Cretaceous of Nălaţ-Vad (Romania), left frontal (UBB NVZ1-38) in dorsal (A) and ventral (B) views. Scale bar: 2 cm. by a narrower hemicylindical neck. On the other anterior to the coronoid process, the lateral surface hand, the condyle is directly connected with a of the dentary body is fl at to slightly convex. It pair of prominent sphenoccipital tubercles, pro- forms a distinct angle with the widely developed jecting lateroventrally from the basioccipital and buccal platform, which separates the coronoid the basisphenoid. Th is is the condition usually process from the dentition and extends anteriorly observed in hadrosaurids. It must be noted that along the full length of the tooth row (Fig. 9A). in Telmatosaurus the occipital condyle is also sup- Weishampel et al. (2003) regarded this important ported by a short but distinct neck (Weishampel development of the buccal platform as one of the et al. 1993). main autapomorphies for Z. shqiperorum. Tw o Between the sphenooccipital tubercles, the ven- or three large neurovascular foramina open along tral side of the basioccipital-basisphenoid complex the margin of the buccal platform anterior to the forms a deep fossa that may mark the attachment coronoid process. site for m. rectus capitis anterioris (Fig. 8A). Th e Th e coronoid process is relatively stout and in- deep carotid canal runs obliquely between the clined posteriorly. It gradually rises from the marked sphenoccipital tubercles and the dorsal part of the angle between the lateral wall of the dentary body broken basipterygoid processes (Fig. 8B). Anteriorly and the buccal platform. Its lateral surface is stri- on the endocranial surface of the basisphenoid, ated, refl ecting the attachment of the adductor the hypophyseal cavity forms a deep, but narrow musculature. Its medial surface is in close contact depression. with the coronoid bone, and its posterior surface with the surangular. Dentary (Fig. 9) (UBB NVZ1-1 and NVZ1-2) In dorsal view, the tooth row is straight. It As previously noted by Weishampel et al. (2003), contains nine tooth positions. In medial view, the dentary of Z. shqiperorum is more massively a robust bony parapet conceals the dentition. A constructed than that of Z. robustus. In lateral view, curved line of alveolar foramina and interconnect- the ventral and dorsal borders of the dentary are ing groove forms the ventral limit of the parapet parallel and regularly curved ventrally. Beneath and (Fig. 9B). Th e adductor fossa is vast and extends

534 GEODIVERSITAS • 2009 • 31 (3) Osteology of Zalmoxes shqiperorum (Dinosauria, Ornithopoda) from Romania

frontal suture A

anterior plate supratemporal fenestra

squamosal orbital margin process anterior plate frontal facet BC

orbital anterior margin plate

laterosphenoid curved ledge jugal process squamosal recess process jugal process

FIG. 7. — Left postorbital (UBB NVZ1-41) of Zalmoxes shqiperorum Weishampel, Jianu, Cziki & Norman, 2003 from the Upper Creta- ceous of Nălaţ-Vad (Romania) in dorsal (A), lateral (B), and medial (C) views. Scale bar: 3 cm.

A floor of foramen magnum

occipital condyle

sphenooccipital fossa for m. rectus tubercle capitis anterior B hypophyseal cavity basioccipital occipital C condyle

carotid canal

basi- occipital sphenooccipital tubercle basisphenoid basisphenoid

sphenooccipital tubercle

FIG. 8. — Basioccipital-basisphenoid complex (UBB NVZ1-40) of Zalmoxes shqiperorum Weishampel, Jianu, Cziki & Norman, 2003 from the Upper Cretaceous of Nălaţ-Vad (Romania) in posterior (A), right lateral (B), and ventral (C) views. Scale bar: 2 cm.

GEODIVERSITAS • 2009 • 31 (3) 535 Godefroit P. et al.

A

coronoid B surangular D buccal platform

coronoid surangular surangular coronoid process coronoid process E C predentary suture

parapet

external mandibular symphysis fenestra external mandibular mandibular canal fenestra

FIG. 9. — Left dentary – surangular – coronoid complex (UBB NVZ1-1) of Zalmoxes shqiperorum Weishampel, Jianu, Cziki & Norman, 2003 from the Upper Cretaceous of Nălaţ-Vad (Romania), in dorsal (A), medial (B, C), and lateral (D, E) views. Scale bar: 5 cm. anteriorly as the mandibular canal nearly the close to the anterior border, towards the adductor length of the dentary. It terminates just posterior fossa, is interpreted by Weishampel et al. (2003) as to the mandibular symphysis. Th e symphysis forms a reduced external mandibular fenestra. an irregular inclined surface. As is usual in orni- thopods, the articular surface for the predentary Predentary (Fig. 10) (UBB NVZ1- 35) forms, when the paired dentaries are articulated, Th e predentary is very high, deep, and subtriangular a scoop-shaped surface. Th ere is no diastema in lateral view. Its anterior border forms an angle of between the last dentary tooth and the articular approximately 45° to the horizontal. Numerous grooves surface for the predentary. and ridges ornament its external surface, suggesting that it was covered by a keratinous rhamphotheca in Coronoid (Fig. 9A, B) (UBB NVZ1-1 and NVZ1-2) life. Unlike in Z. robustus, the oral margin is chisel- Th e coronoid is tightly applied along the medial like, with a particularly sharp outer edge (Fig. 10B). aspect of the coronoid process of the dentary. Th e Th e U-shaped posteroventral surface of the preden- separation between both elements can be observed tary, which articulated with the anterior portion of as a deep groove along the apex of the coronoid the paired dentaries, is formed by a deep and wide process (Fig. 9B). groove, surrounded by high lateral and medial ridges that stabilized the symphyseal region of the mandible Surangular (Fig. 9) (UBB NVZ1-1 and NVZ1-2) (Fig. 10C). Th e high lateroventral processes, regarded Th e surangular is very intimately applied along the by Weishampel et al. (2003) as diagnostic for Zalmoxes, posterior side of the dentary. A small rounded fora- are not preserved in the available material. Th erefore, men that opens on the dorsal part of the surangular, it cannot be decided yet whether this character is really

536 GEODIVERSITAS • 2009 • 31 (3) Osteology of Zalmoxes shqiperorum (Dinosauria, Ornithopoda) from Romania

ABCoral margin oral margin

dentary facet

dentary facet

FIG. 10. — Predentary (UBB NVZ1-35) of Zalmoxes shqiperorum Weishampel, Jianu, Cziki & Norman, 2003, from the Upper Cretaceous of Nălaţ-Vad (Romania), in right lateral (A), dorsal (B), and ventral (C) views. Scale bar: 4 cm.

ABprimary ridge primary C wear facet ridge subsidiary ridges

basal lip

FIG. 11. — Isolated dentary tooth (UBB NVZ1-43a) of Zalmoxes shqiperorum Weishampel, Jianu, Cziki & Norman, 2003 from the Upper Cretaceous of Nălaţ-Vad (Romania), in lingual (A), labial (B), and mesial (C) views. Scale bar: 2 cm. synapomorphic for both Z. robustus and Z. shqipero- enamel is distributed on both sides of the crown. Th e rum, or autapomorphic for Z. robustus. labial side is usually heavily worn (Fig. 11B). Wear begins on the distal half of the crown. A second facet Dentary teeth (Fig. 11) (UBBNVZ1-43) develops mesially as attrition continues. Th e two Th e dentary teeth of Z. shqiperorum are particularly facet merge and cannot be distinguished on heavily- massive. Th e crown is slightly taller than wide. Th e worn teeth. Th e angle of wear ranges between 55

GEODIVERSITAS • 2009 • 31 (3) 537 Godefroit P. et al.

neural spine A neural spine transverse C process transverse process B para- pophysis prezygapophysis parapophysis post- zygapophysis

diapophysis parapophysis

FIG. 12. — Dorsal vertebra (UBB NVZ1-5) of Zalmoxes shqiperorum Weishampel, Jianu, Cziki & Norman, 2003 from the Upper Creta- ceous of Nălaţ-Vad (Romania), in anterior (A), left lateral (B), and posterior (C) views. Scale bar: 4 cm. and 75° to the horizontal. Th e lingual side of the ventrally and the ventral border of the centrum crown is characterized by a strong primary ridge, has consequently an arched aspect (Fig. 12B). Th e slightly mesial to the midline of the crown (contra prezygapophyses face dorsally and medially. Th ey Weishampel et al. 2003). From eight to 13 slightly are better separated from each other on the most divergent vertical subsidiary ridges cover either side anterior vertebrae. On the most anterior vertebrae of the median ridge (Fig. 11A). Th e mesial edge of of the preserved series, the transverse processes are the crown bears tiny crenulations. Th ere is no real oriented posterodorsally; posteriorly in the dorsal cingulum, but a thin enamelled lip marks the base series, the transverse processes become straighter, of the distal part of the crown. Th e root is some- more horizontal and stouter. Th e postzygapophyses what higher than the crown and regularly curved face ventrally and laterally. Th ey increase in size and labially. Grooves on either side of the root and at curvature posteriorly. In lateral view, the neural the base of the crown indicate that the teeth were spine is approximately as high as the centrum and closely packed (Fig. 11C). sub-rectangular in shape. It is more inclined pos- teriorly on the anterior than on the most posterior Dorsal vertebrae (Fig. 12) (UBB NVZ1-3, dorsal vertebrae of the preserved series. NVZ1-4 , NVZ1-5, NVZ1-6, NVZ1-7, NVZ1-12 , NVZ1-13, NVZ1-15, NVZ-7) Sacrum (Fig. 13) (UBB NVZ1-9, NVZ1-10, Th e centrum of the dorsal is approximately as long NVZ1-11, UBB SPZ-2) as high, usually very contracted between the articular UBB SPZ-2 has already been briefl y described and surfaces and keeled along its ventral surface. Th e fi gured by Weishampel et al. (2003). However, it centrum of more posterior dorsal vertebrae becomes appears that there are some minor misinterpretations larger, the lateral sides become more contracted and in their description. Moreover, the preparation of the ventral keel becomes more rounded. Th e articular this specimen has been completed in the meantime. surfaces are slightly amphicoelous to amphiplatyan. Th erefore, it has been decided to include this mate- Th e anterior surface is usually larger than the pos- rial in the description of Z. shqiperorum. Despite terior surface. In lateral view, the planes formed by its fusion to the sacrum, the fi rst vertebra of the the anterior and posterior articular surfaces converge series can be interpreted as the last dorsal vertebra.

538 GEODIVERSITAS • 2009 • 31 (3) Osteology of Zalmoxes shqiperorum (Dinosauria, Ornithopoda) from Romania

ischium last dorsal

s1 s2 s4s3 s5 s6

sacrodorsal A B pubis ilium

pubis ischium last dorsal

s1 C sacrodorsal pubis ilium sacrodorsal D ischium last dorsal

s1 s4 s5 s6

E F

FIG. 13. — Articulated sacrum, ilia, ischia, and left pubis (UBB SPZ-2) of Zalmoxes shqiperorum Weishampel, Jianu, Cziki & Norman, 2003 from the Upper Cretaceous of Nălaţ-Vad (Romania), in dorsal (A, B), left lateral (C, D), and ventral (E, F) views. Scale bar: 10 cm.

Its centrum is sub-circular in cross-section and its the medial surface of the preacetabular process of lateral sides are strongly contracted. In lateral view, the ilium as well. its articular surfaces slightly converge ventrally. Th ere are six fi rmly co-ossifi ed true sacral verte- Th e ventral keel is very rounded on its centrum. brae. Th e centrum of the first one is particularly Its transverse processes are stout. Its neural spine massive, with a posterior articular surface much is rectangular, with a thickened upper edge, and is larger than the anterior surface. Th e centra of the slightly inclined posteriorly. following vertebrae progressively become narrower. Th e second vertebra of the series is somewhat Th e ventral part of the sacral centra forms a shallow larger than the fi rst, but it is very similar in shape. continuous groove, well visible on UBB NVZ1-9 However, it can be regarded as a sacrodorsal vertebra and NVZ1-11. At the junction between adjacent instead of a dorsal vertebra, because the rib of the vertebrae, fi ve massive sacral ribs are fused together succeeding fi rst true sacral vertebra has migrated to form a broad sacrocostal yoke, which attaches anteriorly to fuse across the articulation between to the internal surface of the ilium in the region both adjacent vertebrae. Th is fi rst sacral rib is directly spanning the acetabulum. Th e neural arches of the connected to the medial side of the preacetabular sacral vertebrae slightly overlap the centrum of the process of the ilium and does not participate in the preceding vertebra. Th e prezygapophyses are very sacrocostal yoke. Th e rib of the sacrodorsal vertebra elongated to fi t with the postzygapophyses of the is not preserved, but it may have been attached to preceding sacral vertebra. Th e neural spines of the

GEODIVERSITAS • 2009 • 31 (3) 539 Godefroit P. et al.

A prezygapophysis C postzygapophysis prezygapophysis B

caudal rib

FIG. 14. — Caudal vertebra (UBB NVZ1-14) of Zalmoxes shqiperorum Weishampel, Jianu, Cziki & Norman, 2003 from the Upper Cre- taceous of Nălaţ-Vad (Romania), in anterior (A), dorsal (B), and left lateral (C) views. Scale bar: 4 cm.

sacrals are not well preserved, but they were low, from that of Z. robustus. Th e shaft of the scapular rectangular in lateral view, and their thickened upper blade is very narrow and strap-like. Its posterodorsal edges were at least partially fused together. end sharply expands posteriorly. Th e ventral end of the scapula is also much expanded. Th e anteroventral Caudal vertebrae (Fig. 14) (UBB NVZ1-8, acromial region is fl ared and its lateral surface bears NVZ1-14, NVZ-8, NVZ-15, NVZ-16) a strong deltoid ridge, which limits a vast deltoid Th e articular surfaces of the caudal vertebrae of fossa (Fig. 15A). Both the sutural surface for the Z. shqiperorum are slightly amphicoelous and nearly coracoid and the glenoid part of the scapula are as wide as high. In lateral view, the proximal and thick, moderately concave and very rugose. distal articular surfaces are sub-parallel and inclined posteriorly. Th e distal articular surface is higher Coracoid (Fig. 15D, E) (UBB NVZ1) than the proximal surface and the ventral margin, Th e coracoid is characterized by its unusually long between both articular surfaces, is strongly concave. and prominent posterior sternal process, as previ- Th e haemapophyseal facets are more distinct on the ously described by Weishampel et al. (2003). Th e ventral surface of UBB NVZ1-8 than in the more glenoid facet is reniform and concave at its centre. proximal vertebrae. A pair of longitudinal ridges Th e glenoid facet forms a prominent lip on the lat- joins the haemapophyseal facets and the ventral eral surface of the coracoid. Th e coracoid foramen surface of the centrum looks consequently furrowed. perforates the thickest part of the bone, beneath Th e slender caudal ribs are fused to the sides of the the scapulocoracoid suture. centra along the neurocentral suture. Th e pre- and postzygapophyseal processes become more elongated Humerus (Fig. 16) (UBB NVZ2-3, NVZ2-4, as the size of the neural arch diminishes and as the NVZ-5) centrum becomes proportionally more elongated, and Th e humerus of Z. shqiperorum diff ers from that the zygapophyseal surfaces become nearly vertical. of Z. robustus by the important development of its deltopectoral crest, which extends over the proximal Scapula (Fig. 15A-C) (UBB NVZ1-21, NVZ1-24, end of the humerus. Th e edge of the deltopecto- NVZ2-2) ral crest is regularly concave and turned medially, As previously described by Weishampel et al. (2003), and its distal tip is very salient. Th e deltopectoral the scapula of Z. shqiperorum is markedly diff erent crest therefore limits a wide bicipital sulcus on the

540 GEODIVERSITAS • 2009 • 31 (3) Osteology of Zalmoxes shqiperorum (Dinosauria, Ornithopoda) from Romania

A B C glenoid D

sternal D, E process glenoid E

deltoid fossa coracoid foramen deltoid sternal glenoid ridge process

A-C coracoid facet acromion

FIG. 15. — Zalmoxes shqiperorum Weishampel, Jianu, Cziki & Norman, 2003 from the Upper Cretaceous of Nălaţ-Vad (Romania), right scapula (UBB NVZ1-24) in lateral (A), medial (B), and dorsal (C) views; left coracoid (NVZ1-36) in medial (D) and lateral (E) views. Scale bars: A-C, 5 cm; D, E, 4 cm. anterior surface of the humerus (Fig. 16A). Th e triangular crest that progressively merges with the globular proximal articular head forms a rounded ulnar shaft. On the proximodorsal surface of the buttress on the posterior side of the humerus. Th e ulna, the articular facet for the proximal part of inner tuberosity is well developed and forms a the radius is large, triangular in shape and concave. buttress beneath the concave medial border of the Th e ulna progressively tapers distally. Its distal end humerus (Fig. 16D). Because of the important is mediolaterally compressed. development of the deltopectoral crest and of the distal portion of the humerus, the humeral shaft Ilium (Figs 13; 18A-D) (UBB NVZ1-16, looks proportionally slender. Th e distal portion NVZ1-17, UBB SPZ-2) of the humerus is mediolaterally expanded and Th e ilium of Z. shqiperorum is particularly massive somewhat off set medially. Th e ulnar condyle is in dorsal view, because the lateral side of the iliac much larger and extends more distally than the body faces dorsomedially, forming an angle of about radial condyle. Th e olecranon fossa forms a broad 30° to the horizontal plane. Moreover, the dorsal triangular depressed area on the posterior surface edge of the ilium is sigmoidal and particularly of the humerus (Fig. 16B). Th e coronoid fossa is thick (Fig. 18A). It is ornamented by numerous poorly developed (Fig. 16A). longitudinal ridges, indicating that it served as at- tachment site for a strong pelvic musculature (see Ulna (Fig. 17) (UBB NVZ-23) e.g., Galton 1974 and Norman 1986, for a detailed Th e ulna of Z. shqiperorum is very robust and description of the musculature of the pelvic girdle slightly curved medially. Th e olecranon process is and hind limb in ornithopods). Th e preacetabular prominent, but short and blunt. Th e proximomedial process is long and dorsoventrally fl attened. It is coronoid process is developed into a prominent curved laterally and its dorsal surface slightly faces

GEODIVERSITAS • 2009 • 31 (3) 541 Godefroit P. et al.

humeral head AB

delto- pectoral inner tuberosity crest humeral head C D bicipital sulcus

deltopectoral crest

coronoid fossa ulnar condyle olecranon fossa

FIG. 16. — Zalmoxes shqiperorum Weishampel, Jianu, Cziki & Norman, 2003 from the Upper Cretaceous of Nălaţ-Vad (Romania), left humerus (UBB NVZ4) in anterior (A) and posterior (B) views; left humerus (UBB NVZ5) in anterior (C) and posterior (D) views. Scale bars: 5 cm. laterally. A prominent longitudinal groove, limited is everted to form a prominent supra-acetabular dorsally by a high and sharp ridge, runs along its shelf (Fig. 18A), much more developed than in medial surface (Fig. 18B). Th e preacetabular notch Z. robustus. Th e postacetabular process is long is very deep. Above the acetabulum, the body of the and robust. Its dorsal margin much more everted ilium is lower than in Z. robustus. Th e morphology laterally than in Z. robustus, in continuity with the of the acetabulum is fundamentally diff erent between supra-acetabular shelf. both species. In Z. robustus, the acetabulum is of limited extend, forming a shallow breach between Ischium (Figs 13; 18E, F) (UBB NVZ1-18, the enormous, lenticular ischial peduncle and the NVZ1-21, NVZ1-22, UBB SPZ-2) smaller pubic peduncle. In Z. shqiperorum, on the Th e iliac peduncle is much more developed in other hand, the pubic peduncle is better developed, Z. shqiperorum than in Z. robustus and forms a extending below the level of the ischial peduncle, slightly depressed elliptical surface that fi ts the and the acetabulum forms a well-developed and globular ischial peduncle of the ilium. Th e pubic regularly concave embayment on the ventral part peduncle is more slender and elongate. Between the of the ilium (Fig. 18A). Th e acetabular surface is iliac and pubic peduncles, a shallow embayment limited laterally, from the most anterior point of forms the posteroventral margin of the acetabu- the ischial penduncle to the tip of the pubic pedun- lum. Th e ischial shaft is mediolaterally attened.fl cle, by a continuous ridge. Above and posteriorly Its distal half is curved ventrally. Th e distal end of to the acetabulum, the dorsal margin of the ilium the ischium is curved laterally and forms a promi-

542 GEODIVERSITAS • 2009 • 31 (3) Osteology of Zalmoxes shqiperorum (Dinosauria, Ornithopoda) from Romania

nent boot-like expansion. Its medial side is fl at olecranon process olecranon process and bears numerous striations, indicating a strong proximomedial ligamentous attachment with the paired element. coronoid Like in Z. robustus, there is no obturator process A process B on the ischial shaft. proximal Pubis (Fig. 13) (UBB SPZ-2) radius Both the iliac and the ischial peduncles are wide facet and cup-shaped. Th e small embayment between them forms the anteroventral margin of the acetab- ulum. Th ere is apparently no pubic foramen. Th e prepubic neck is long and very slender. Th e prepubis is also very slender and only slightly expanded dorsally.

Femur (Fig. 19) (UBB NVZ1-25, NVZ-1, NVZ2-5, NVZ2-6, NVZ-9, NVZ-21) Th e femur of Z. shqiperorum closely resembles that of Z. robustus in being distinctly bowed laterally in anterior view and nearly straight in lateral view. Th e femoral head is massive and hemispherical. It is set off from the long axis of the shaft by a rela- tively long neck. Th e greater trochanter is regularly rounded and is somewhat lower than the femoral head. Th e anterior trochanter is apparently bet- ter developed than in Z. robustus, particularly in larger specimens. It is separated from the greater trochanter by a vertical cleft. Its external surface FIG. 17. — Right ulna (UBB NVZ23) of Zalmoxes shqiperorum Weishampel, Jianu, Cziki & Norman, 2003 from the Upper Creta- bears strong striations, marking the insertion area ceous of Nălaţ-Vad (Romania), in anterior (A) and medial (B) views. for a powerful m. ilio-femoralis externus. Th e lateral Scale bar: 5 cm. side of the greater trochanter forms an extended depressed area, limited anteriorly by the anterior trochanter, and posteriorly and ventrally, by a a prominent internal condyle that overhangs the strong but smooth ridge (Fig. 19D). Its striated tibial shaft. It is separated by a shallow popliteal surface served as attachment site for a very pow- groove from the smaller lateral condyle. Th e cnemial erful m. ilio-trochantericus. Th e pendant fourth crest is less prominent than in Z. robustus and more trochanter is placed at the mid-shaft. Th e medial derived ornithopods, and occupies only about the condyle is stouter, but the lateral condyle extends proximal fi fth of the tibia. Its anteroproximal surface further distally. Th e fl exor intercondylar groove is forms a deep and wide anterior groove, probably much deeper and wider than the extensor inter- marking the attachment area for the strong distal condylar groove. Th e posterolateral surface of the ligament of m. ilio-tibialis (Fig. 20B). Th is groove lateral condyle forms a long but shallow vertical has not been described in Z. robustus (Weishampel groove for m. iliofi bularis (Fig. 19D). et al. 2003). Th e tibial shaft is long, more slender than in Z. robustus, and ovoid in cross-section. Tibia (Fig. 20A-D) (UBB NVZ-2, NVZ-3) In anterior view, it is less bowed laterally than in Th e tibia is less massive in Z. shqiperorum than Z. robustus. Below the end of the cnemial crest, a in Z. robustus. Its posteroproximal angle forms round vertical ridge descends to the lateral edge

GEODIVERSITAS • 2009 • 31 (3) 543 Godefroit P. et al.

supra-acetabular shelf preacetabular process A

postacetabular process preacetabular notch

ischial peduncle pubic peduncle acetabulum preacetabular process B A, B

postacetabular process preacetabular notch

C acetabulum pubic peduncle

supra-acetabular shelf

D preacetabular process

preacetabular notch C, D pubic peduncle

iliac peduncle ischial shaft ischial shaft E F

acetabulum

pubic peduncle

FIG. 18. — Zalmoxes shqiperorum Weishampel, Jianu, Cziki & Norman, 2003 from the Upper Cretaceous of Nălaţ-Vad (Romania), left ilium (UBB NVZ1-17) in lateral (A) and medial (B) views; right ilium (UBB NVZ1-16) in dorsal (C) and lateral (D) views; right ischium (UBB NVZ1-22) in lateral view (E); left ischium (UBB NVZ1-18) in medial view (F). Scale bars: 5 cm. of the lateral malleolus. On the distal end of the Fibula (Fig. 20E, F) (UBB NVZ1-28) tibia, the external malleolus is very salient distally. Like in Z. robustus, the fi bula of Z. shqiperorum is Th e internal malleolus is, on the other hand, more slender. Th e medial surface of the proximal half of salient medially, with an articular surface turned the shaft forms a concave surface that is irregularly towards the anterolateral side of the tibia. bordered by smooth anterior and posterior ridges.

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insertion area for m. ilio-femoralis externus greater trochanter greater trochanter anterior femoral head femoral head ABCDtrochanter

anterior troch- anter

4th trochanter

medial condyle lateral condyle

medial condyle flexor groove groove for m. iliofibularis lateral extensor groove condyle

FIG. 19. — Right femur (UBB NVZ9) of Zalmoxes shqiperorum Weishampel, Jianu, Cziki & Norman, 2003 from the Upper Cretaceous of Nălaţ-Vad (Romania), in anterior (A), medial (B), posterior (C), and lateral (D) views. Scale bar: 5 cm.

Th is surface is also ornamented by longitudinal the distal end. Its medial side is smoothly convex, ridges, indicating that it was fi rmly attached to the whereas its lateral side is fl at. Th e distal articular tibial shaft by ligaments. surface is regularly convex and it slightly faces me- dially. A shallow intercondylar groove is developed Metatarsals on its plantar aspect and probably acted as a pulley Metatarsal II of Z. shqiperorum (UBB NVZ1-30; to guide a powerful fl exor tendon. Fig. 20G, H) is elongated and rather slender. Its Only the distal portion of metatarsal III (UBB proximal end is mediolaterally compressed and NVZ1-34; Fig. 20I, J) is preserved. It is rather slender plantodorsally expanded. Its medial surface bears too, with a trapezoidal dorsoplantarly fl attened shaft, strong longitudinal striations, indicating ligamen- whose medial side is higher than its lateral side. Th e tous attachment with metatarsal I. Its lateral side distal end of metatarsal III is expanded into a large forms a fl at triangular surface that also bears lon- saddle-shaped and dorsoplantarly convex articular gitudinal striations marking strong attachment to surface. Th e intercondylar groove is well developed metatarsal III. Th e distal end of metatarsal II is both on the dorsal and plantar parts on the condyle. plantodorsally expanded and only a little wider than Th e medial condyle is more developed than the lateral

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condyle. Both the medial and lateral sides of the distal re-examination of UBB SPZ-2 and discovery of articular end form cup-like depressions. new material at Nălaţ-Vad reveal that the supra- acetabular process is well developed in Z. shqipero- rum as well. On the other hand, it is apparently SYSTEMATIC OBSERVATIONS not developed in R. priscus (Pincemaille-Quillévéré 2002: fi g. 15) and other basal Iguanodontia (Nor- Weishampel et al. (2003) based their description man 2004). Th erefore, a well-developed supra- of Z. shqiperorum mainly on postcranial mate- acetabular process can be regarded as a diagnostic rial, collected in various localities in the Haţeg character for Zalmoxes. Basin. Of the cranial material, only the dentary of Weishampel et al. (2003) considered that a re- Z. shqiperorum was known at that time. Th e dis- duced external mandibular fenestra, positioned covery at Nălaţ-Vad of fairly well-preserved skull along the upper anterior border of the surangular material, in association with postcranial elements, and overlapped by the dentary, and the develop- is therefore important for a better understanding ment of a secondary surangular foramen are also of this species and, more generally, of Zalmoxes. synapomorphic for Zalmoxes. Th e description of the Indeed, in their diagnosis of Zalmoxes, Weisham- material from Nălaţ-Vad confi rms that Z. shqipero- pel et al. (2003) listed a number of characters that rum is characterized, like Z. robustus, by a reduced were in fact not known in the material pertaining external mandibular fenestra (the posterior part of to Z. shqiperorum or in the diff erent species of the surangular, with the secondary foramen, is not Rhabdodon. Consequently it was in fact impossi- preserved). However, Pincemaille-Quillévéré (2002) ble to decide whether these characters were really also describes two foramina on the external surface synapomorphic for Zalmoxes, synapomorphic for of the surangular in R. priscus. It is therefore quite Rhabdodontidae, or autapomorphic for Z. robustus. possible that those foramina respectively repre- Th e study of the Nălaţ-Vad material confi rms that sent a reduced external mandibular fenestra and a the following characters can be observed in both secondary mandibular foramen. In this case, both Z. robustus and Z. shqiperorum: characters would represent synapomorphies for the 1. extensive, complex squamose suture between Rhabdodontidae clade. In any case, the reduced ex- quadratojugal and jugal; ternal mandibular fenestra is placed more anteriorly 2. post-temporal foramen transmitted through the in Z. robustus and Z. shqiperorum than the anterior body of the squamosal; foramen on the surangular of R. priscus. 3. curved shelf on the lateral surface of the pos- Th e following characters, regarded as diagnostic torbital; for Zalmoxes by Weishampel et al. (2003), cannot be 4. lateral splaying of the quadrate; observed either in Z. shqiperorum or in the diff erent 5. deep predentary; species of Rhabdodon; therefore, subsequent discover- 6. long, dorsoventrally-narrow, twisted pre acetabular ies may shift their distribution either inclusively (as process; synapomorphies for the Rhabdodontidae clade) or 7. absence of an obturator process on the ischium; exclusively (as autapomorphies for Z. robustus): 8. arched ischial shaft. 1. absence of the scar for m. adductor mandibulae Nevertheless, only the last two characters can be externus superfi cialis on the squamosal; regarded as strictly synapomorphic for Zalmoxes. 2. a large, disc-shaped quadratojugal; Indeed, the state of the other features remains un- 3. a frontal with a complex transverse sutural sur- known in Rhabdodon and it cannot therefore be face that extensively overlaps the parietal; excluded that they represent in fact synapomorphies 4. high lateroventral processes on the predentary. for Rhabdodontidae. Th e following characters appear autapomorphic Weishampel et al. (2003) regarded the presence for Z. shqiperorum. Indeed, they have not been of a well-developed supra-acetabular process on the observed either in Z. robustus or in other basal ilium as an autapomorphy for Z. robustus. However, Iguanodontia:

546 GEODIVERSITAS • 2009 • 31 (3) Osteology of Zalmoxes shqiperorum (Dinosauria, Ornithopoda) from Romania

propliteal groove lateral condyle internal condyle anterior groove cnemial internal condyle cnemial crest A B crest

C D

internal internal malleolus external external internal external malleolus malleolus malleolus malleolus malleolus EF proximal tibial facet G I

J H

FIG. 20. — Zalmoxes shqiperorum Weishampel, Jianu, Cziki & Norman, 2003 from the Upper Cretaceous of Nălaţ-Vad (Romania), left tibia (UBB NVZ3) in lateral (A), anterior (B), medial (C), and posterior (D) views; ?left fi bula (UBB NVZ1-28) in lateral (E) and medial (F) views; left metatarsal II (UBB NVZ1-30) in medial (G) and lateral (H) views; ?left metatarsal III (UBB NVZ1-34) in plantar (I) and ?medial (J) views. Scale bars: A-D, 5 cm; E, F, 4 cm; G-J, 3 cm.

GEODIVERSITAS • 2009 • 31 (3) 547 Godefroit P. et al.

1. the occipital condyle is not separated from the suffi cient to support the validity of both species. sphenooccipital tubercles by a distinct neck; Th ere is no indication that these diff erences refl ect 2. the dentary bears an angular buccal emargination ontogenetic variability within a single species: smaller that forms a horizontal platform extending for the and larger individuals are clearly represented in full length of the dentition behind and medial to both groups. Th e diff erences between Z. robustus the coronoid process (Weishampel et al. 2003); and Z. shqiperorum also appear too important to 3. the scapular blade is narrow and strap-like proxi- simply refl ect sexual dimorphism. As observed in mally and expands sharply posterodistally (Wei- lambeosaurines (Dodson 1975), in basal necera- shampel et al. 2003); topsians (Dodson 1976), or in ceratopsids (Lehman 4. the region of the scapula adjacent to the cora- 1990; Sampson et al. 1997), dinosaur males and coid suture is expanded; acromial process forming females may not be distinguished by the presence a prominent fl ange (Weishampel et al. 2003); or absence of osteological characters, but instead 5. the deltopectoral crest of the humerus is par- by the diff erence in development of cranial super- ticularly prominent, extending over the proximal structures (crests, horns, frills) at a given size. It is half of the humerus; now usually admitted that these structures primarily 6. the iliac peduncle of the ischium is particularly functioned as signals to recognize and to compete developed; for mate or territory. Such characters are subject 7. the distal end of the ischium forms a boot-like to delayed growth, developing only after the onset expansion (Weishampel et al. 2003). of adult size (Dodson 1975; Sampson et al. 1997). Th e description of the Nălaţ-Vad material reveals Sexual dimorphism in the postcranial skeleton has that Z. shqiperorum potentially diff ers from Z. ro- not been well studied yet. However, Raath (1990) bustus by a series of other characters, listed below. tentatively explained the morphological variation However, these characters cannot be adequately in the robustness of the postcranial skeleton of the quantifi ed in the current state of our knowledge, basal theropod Syntarsus by sexual dimorphism: he because we lack more precise information about suggested that the most robust morph might repre- their intraspecifi c variability. Moreover, their state sent females. In Zalmoxes, important diff erences in remains ambiguous: the skull element and in the postcranium between 1. the jugal appears more gracile in Z. shqiperorum; the two supposed species are not simple variations in 2. the parietal process of the squamosal is appar- the development of supracranial structures or in the ently shorter, but higher; robustness of the bones, but true osteological. For 3. the infratemporal notch on the squamosal ap- examples, it seems diffi cult to explain the observed pears lower, but deeper; diff erences in the morphology of the basioccipital, 4. the post-temporal foramen is set more ante- the scapula or the ischium by sexual dimorphism: riorly; these characters cannot be recognized as visual 5. the sagittal crest is better developed; signals. Moreover, these features were apparently 6. the dentary is more massively constructed; already developed in juveniles, before individuals 7. the oral margin of the predentary is apparently have reached sexual maturity. characterized by a sharper outer margin; Figure 1 represents the geographic distribution 8. the anterior trochanter is better developed on of Z. robustus and Z. shqiperorum. Both species the femur; have been discovered together in several localities 9. the tibia appears less massive. (Sănpetru, Tustea, Valioara, Vurpar) and, so far, there is no indication that they were found in dif- ferent levels. Although they clearly occupied very SYMPATRIC SPECIES IN ORNITHOPODS similar ecological niches and probably competed for the same food resources, the two species there- Th e observed morphological diff erences between fore appear sympatric. As discussed below, it is not Z. robustus and Z. shqiperorum appear, in any case, the fi rst time that two closely-related species of an

548 GEODIVERSITAS • 2009 • 31 (3) Osteology of Zalmoxes shqiperorum (Dinosauria, Ornithopoda) from Romania

ornithopod genus are described in a same locality by Norman & Weishampel [1990] and Norman and are therefore regarded as sympatric. [2004]), in the Tilgate Forest area, the historical At Bernissart (Belgium, Early Cretaceous), two place where the fi rst dinosaur specimens were dis- or three specimens of the gracile species Iguanodon covered in the 19th Century (Benton & Spencer atherfi eldensis Hooley, 1925 have been discovered 1995; Weishampel et al. 2004). Unfortunately, together with more than 20 specimens of the robust because the quarries have been closed for a long I. bernissartensis Boulenger, 1881 (Norman 1986). time, most of the discovery sites remain uncer- Paul (2007) proposed that I. bernissartensis and tain. Th e Hastings Beds (Berriasian-Vallanginian) I. atherfi eldensis do not form a monophyletic group that outcrop along the coast east of Hastings (East and that morphological diff erences are suffi cient Sussex, United Kingdom) are also famous for their to include the latter species into a separate genus, vertebrate remains. Th ree Iguanodon species have Mantellisaurus Paul, 2007. However, he conducted been described from this area: I. dawsoni Lydekker, no phylogenetic analysis proving that I. bernissartensis 1888 and I. fi ttoni Lydekker, 1889, from Shorden and I. atherfi eldensis are really paraphyletic. On the Quarry, and I. hollingtonensis Lydekker, 1889, from other hand, in the most recent phylogenetic analysis Hollington Quarry. Norman & Weishampel (1990) of the group, Norman (2004: fi g. 19.22) suggested and Norman (2004) synonymised I. hollingtonensis that both species may be regarded as monophyletic with I. fi ttoni and accepted I. dawsoni as valid, but and, if this hypothesis is correct, it is certainly not do not elucidate the diff erences of these poorly necessary to split them into diff erent genera. In known taxa from the typical I. bernissartensis and any case, it is defi nitely not proved that I. bernis- I. atherfi eldensis. In France, Iguanodon specimens sartensis and I. atherfi eldensis really lived together were found in Early Cretaceous deposits from the in the Bernissart area. Indeed, the Bernissart fossil Saint-Dizier region (Haute-Marne). However, locality is apparently an attritional accumulation the two species were found in diff erent localities: of animals dead in diff erent places and at diff erent I. bernissartensis at Baudonvilliers, Pont-Varin and times. During Early Cretaceous times, the Bernissart Saint-Dizier, and I. atherfi eldensis at Wassy and site may have been somewhat lower topographi- Coursancelles (Martin & Buff etaut 1992; É. Buf- cally than the surrounding countryside because fetaut pers. comm. 2006). Adult and numerous of subterranean dissolution, which should explain juvenile remains of both I. atherfi eldensis and I. ber- the preferential accumulation of fossils at Bernis- nissartensis were discovered in a cavern-like deposit sart (Delmer & Van Wichelen 1980). However, at a quarry near the village of Nehden in Sauerland these two species have also been described together (Germany). Contrary to the Bernissart assemblage, in many other Early Cretaceous formations from the Nehden fauna is dominated by I. atherfi eldensis Western Europe, suggesting that they were really specimens and by juveniles. According to Norman sympatric in this area. Although the holotype of (1987) and Norman et al. (1987), the assemblage I. atherfi eldensis Hooley, 1825 (BMNH R5764) was at Nehden may represent a catastrophic accumula- collected from the Vectis Formation, most of the tion caused by a fl ash fl ood or by a herd crossing a Iguanodon specimens collected in the south-west river. However, the over-representation of juvenile coast of the Isle of Wight, including partial skeletons specimens in a fossil assemblage is a good indicator and isolated bones of both I. atherfi eldensis and for an attritional death profi le (Lyman 1994): the I. bernissartensis, were found in diff erent points of observed peaks corresponding to ages where mor- the underlying Wessex Formation (Martill & Naish tality rates are the highest, among the very young 2001). Th e Smokejacks Pit locality (Surrey, UK) and, to a lesser extent, the very old. Th erefore, it has also yielded good material belonging to both cannot be excluded that the Nehden assemblage I. atherfi eldensis and I. bernissartensis (Benton & represents, like Bernissart, an allochtonous accu- Spencer 1995). Both species were also reported mulation of animals dead in diff erent places and from the Hastings Beds and Weald Clay of West times. Fragmentary remains identifi ed as I. bernis- Sussex, with I. anglicus Holl, 1829 (regarded as valid sartensis and I. atherfi eldensis (“I. mantelli”) have

GEODIVERSITAS • 2009 • 31 (3) 549 Godefroit P. et al.

been described together in the Camarillas Forma- Ariège and Gard in Southern France. Rhabdodon tion (Barremian) of the San Cristobal and Santa septimanicus is currently known by a single right Barbara localities (Teruel Province, Spain; Sanz dentary, from the “Grès à Reptiles” formation in et al. 1984a, b). However, the described material Montouliers (Hérault). Two species of Rhabdodon appears poorly diagnosed and the identifi cation is therefore apparently coexist in the “Grès à Reptiles” mainly based on the size of the bones. Moreover, formation of the Saint-Chinian area in Hérault. Ruiz-Omeñaca & Canudo (2004) consider that Although R. septimanicus was doubted by some the material from Teruel Province previously re- authors (Allain & Pereda-Suberbiola 2003), it was ferred to as I. bernissartensis would in fact belong accepted by Weishampel et al. (2003) and Norman to a new iguanodontian genus. In conclusion, the (2004). In any case, Buff etaut (2005) indicated that distribution of the diff erent Iguanodon species the abundant Rhabdodon material from the Saint- clearly shows the diffi culty to decide whether the Chinian area apparently includes both a robust and presence of two species within a same formation a gracile form. A detailed study of Rhabdodon mate- results from the sympatry of these species or from rial is currently in progress, which should provide taphonomic processes. a fi nal answer to the question of how many species Th e famous Quarry 13 in the Morrison Formation of Rhabdodon are present in the Late Cretaceous near Como (Wyoming) has yielded the holotypes of of southern France (É. Buff etaut and P. Chanthasit four Camptosaurus Marsh, 1885 species: C. dispar pers. comm. November 2006). (Marsh, 1879), C. medius Marsh, 1894, C. nanus Marsh, 1894 and C. browni Gilmore, 1909. Nor- man & Weishampel (1990), and Norman (2004) CONCLUSIONS considered these taxa as synonyms, but regarded C. amplus (Marsh, 1879), from the neighbouring Th e new fossils discovered in the Sănpetru Formation Quarry 1A at Como Bluff , as a valid species. How- at Nălaţ-Vad include the most complete skeleton ever, this latter taxon is based on a pes, which closely that can be referred to date to Zalmoxes shqiperorum. resembles that of theropod dinosaurs. Th e observed morphological diff erences between Th e Dinosaur Park Formation of Alberta (Canada) Z. robustus and Z. shqiperorum cannot be adequately has yielded at least eight valid hadrosaurid species explained by ontogenetic variation or sexual di- (Ryan & Evans 2004), including two valid species morphism. Because Z. robustus and Z. shqiperorum of Lambeosaurus Parks, 1923 (L. lambei Parks, were discovered together in several localities, their 1923 and L. magnicristatus Sternberg, 1935) and sympatry is therefore a hypothesis that cannot be two species referred to Gryposaurus Lambe, 1914 a priori rejected. Th e sympatry of closely-related (G. notabilis Lambe, 1914 and G. incurvimanus species was apparently not an isolated case among Parks, 1920). Recent data on the stratigraphic ornithopod dinosaurs, even if it is not always easy position of articulated and associated skeletons to decide whether the presence of species belonging within the Dinosaur Park Formation indicate that to a same genus within a same locality is the result Lambeosaurus magnicristatus has no stratigraphic of true sympatry, of taphonomic processes, or of overlap with L. lambei and suggests that it replaces imprecision in the collecting information. L. lambei on a regional scale in southern Alberta (Evans & Reisz 2007). Currie & Russell (2004) also indicate that all Gryposaurus incurvimanus specimens Acknowledgements are higher in the Dinosaur Park Formation than We wish to thank all the participants to the 2002 specimens of G. notabilis. fi eld campaign at Nălaţ-Vad: Paul Dica, Cristina Two species are also currently recognized within Fărcas, Emmanoil and Liana Săsăran, Th ierry Smith, Rhabdodon, the sister-taxon of Zalmoxes. Rhabdodon Pierre Bultynck, Annelise Folie, Jimmy Van Itter- priscus is recorded in various formations and locali- beeck, Suzanne Clinet, Dan Grigorescu and Zoltan ties from Aude, Hérault, Bouches-du-Rhônes, Var, Cziki. Th e fi eldwork was supported by a research

550 GEODIVERSITAS • 2009 • 31 (3) Osteology of Zalmoxes shqiperorum (Dinosauria, Ornithopoda) from Romania

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Submitted on 5 October 2007; accepted on 23 December 2008.

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