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Hadrosauroid dinosaurs from the latest Cretaceous ofthe Iberian PeninsulaXabier Pereda-Suberbiola a , José Ignacio Canudo b , Julio Company c , Penélope Cruzado-Caballero e & José Ignacio Ruiz-Omeñaca b da Universidad del País Vasco/EHU, Facultad de Ciencia y Tecnología, Dpto. Estratigrafía yPaleontología, Apdo. 644, 48080, Bilbao, Spain E-mail:b Grupo Aragosaurus, Paleontología, Facultad de Ciencias, Universidad de Zaragoza, PedroCerbuna 12, 50009, Zaragoza, Spain E-mail: www.aragosaurus.comc Departamento de Ingeniería del Terreno, Universidad Politécnica de Valencia, and Instituto“Cavanilles” de Biodiversidad y Biología Evolutiva, 46022 Valencia, Apartado de Correos22085, 46071, Valencia, Spain E-mail:d Museo del Jurásico de Asturias (MUJA), 33328, Colunga, Asturias, Spain E-mail:e Grupo Aragosaurus, Paleontología, Facultad de Ciencias, Universidad de Zaragoza, PedroCerbuna 12, 50009, Zaragoza, Spain E-mail: www.aragosaurus.com

Version of record first published: 02 Aug 2010

To cite this article: Xabier Pereda-Suberbiola, José Ignacio Canudo, Julio Company, Penélope Cruzado-Caballero & JoséIgnacio Ruiz-Omeñaca (2009): Hadrosauroid dinosaurs from the latest Cretaceous of the Iberian Peninsula, Journal ofVertebrate Paleontology, 29:3, 946-951

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HADROSAUROID DINOSAURS FROM THE LATEST CRETACEOUSOF THE IBERIAN PENINSULA

XABIER PEREDA-SUBERBIOLA,*,1 JOSE IGNACIO CANUDO,2 JULIO COMPANY,3 PENELOPECRUZADO-CABALLERO,2 and JOSE IGNACIO RUIZ-OMENACA2,4; 1Universidad del Paıs Vasco/EHU,Facultad de Ciencia y Tecnologıa, Dpto. Estratigrafıa y Paleontologıa, Apdo. 644, 48080 Bilbao, Spain,xabier.pereda@ehu.es; 2Grupo Aragosaurus (www.aragosaurus.com), Paleontologıa, Facultad de Ciencias,Universidad de Zaragoza, Pedro Cerbuna 12, 50009 Zaragoza, Spain, jicanudo@unizar.es, penelope@unizar.es;3Departamento de Ingenierıa del Terreno, Universidad Politecnica de Valencia, 46022 Valencia, and Instituto“Cavanilles” de Biodiversidad y Biologıa Evolutiva, Apartado de Correos 22085, 46071 Valencia, Spain, company@uv.es;4Museo del Jurasico de Asturias (MUJA), 33328 Colunga, Asturias, Spain, jigruiz@unizar.es

Hadrosauroids were the dominant dinosaurs of the Laurasianlandmasses during the latest Cretaceous (Horner et al., 2004). Themajority of the fossil remains, including fully articulated skeletonswith complete skulls, come from North America and Asia. InEurope, the fossil record is mainly based on disarticulated andisolated material (Dalla Vecchia, 2006). Of nearly 50 localitiesknown in Europe, more than half are in the Iberian Peninsula(Casanovas et al., 1999a; Pereda-Suberbiola et al., 2003; Company,2004; Vila et al., 2006). Most of the sites are Maastrichtian in age,but some remains have also been found in upper Campanian de-posits (Lopez-Martınez et al., 2001; Pereda-Suberbiola et al.,2003). Until recently, only two genera and species had been recog-nized in the Maastrichtian of Europe: Telmatosaurus transsylvani-cus (Nopcsa, 1900) from Transylvania (Weishampel et al., 1993),and Pararhabdodon isonensis Casanovas, Santafe and Isidro, 1993from Catalonia (Casanovas et al., 1999a; Prieto-Marquez et al.,2006). Orthomerus dolloi Seeley, 1883 from the Maastricht areaof the Dutch Limburg is regarded as a nomen dubium (Brink-mann, 1988; Horner et al., 2004). Recently, Koutalisaurus kohler-orum Prieto-Marquez, Gaete, Rivas, Galobart and Boada, 2006has been erected on the basis of a right dentary previously ascribedto P. isonensis (Casanovas et al., 1999a; Pereda-Suberbiola et al.,2003). In addition, hadrosauroid bones and teeth have beencollected during recent years in several Maastrichtian sites ofSpain, mainly in Huesca, Lleida and Valencia provinces (Fig. 1A)(Company et al., 1998; Casanovas et al., 1999a, 1999b; Lopez-Martınez et al., 2001; Company, 2004; Cruzado-Caballero et al.,2005; Vila et al., 2006; Gaete et al., 2007;Martın et al., 2007; Prieto-Marquez et al., 2007). Even though only a portion of this materialis available for study (numerous remains are still unprepared),several taxa seem to be represented. The aim of this paper is toassess the diversity of latest Cretaceous Iberian hadrosauroids andto discuss their paleobiogeographical implications.

METHODS

We follow here the definition of Hadrosauridae outlined byWeishampel et al. (1993) and slightly amended by Horner et al.(2004): the clade consisting of the most recent common ancestorof Telmatosaurus and Parasaurolophus and all their descen-dants. Other works (Forster, 1997; Sereno, 1998; Godefroitet al., 1998; You et al., 2003; Prieto-Marquez et al., 2006) haveargued for a more restrictive definition of Hadrosauridae:Hadrosaurinae plus Lambeosaurinae and their most recent com-mon ancestor (Euhadrosauria of Weishampel et al., 1993).Hadrosauroids are regarded in this paper as all iguanodontiansmore closely related to Parasaurolophus than to Iguanodon

(sensu Sereno, 1986, 1998; You et al., 2003). As noted by Horneret al. (2004), these definitions differ in their membership, but donot alter the phylogenetic relationships of the respective taxa.Institutional Abbreviations—IPS, Institut Catala de Paleonto-

logia (formerly Institut de Paleontologia Dr. Miquel Crusafont),Sabadell, Spain; MCD, Museu de la Conca Della, Isona, Lleida,Spain; MGUV, Museo de Geologıa, Universidad de Valencia,Spain; MPV, Museo Paleontologico de Valencia, Spain; MPZ,Museo Paleontologico, Universidad de Zaragoza, Spain.

THE IBERIAN RECORD

- Pararhabdodon isonensis, Sant Roma d’Abella (Lleida). Thistaxon is known from two maxillae, several vertebrae and limbbones (IPS SRA 1, 12–13, 15–18, 20, 22–26, 693-12, 693-13, MCD4730, 4731) from Sant Roma d’Abella, near Isona (Tremp Basin,Lleida). All the fossils were found in a small surface and presum-ably belong to a single individual (Casanovas et al., 1999a; J.V.Santafe, pers. comm., 1999). Pararhabdodon was originally de-scribed as an iguanodontian close to Rhabdodon, and laterregarded as a basal lambeosaurine (Casanovas et al., 1993,1999a). Recently, Prieto-Marquez et al. (2006) have interpretedPararhabdodon as a basal hadrosaurid (see also Head, 2001) andsister taxon to Euhadrosauria. They redescribed the material andrediagnosed P. isonensis by a few characters of the maxilla (i.e.,highly elevated jugal joint, concave subvertical surface ventral tothe jugal point, anteroposteriorly broad dorsal process) and thehumerus (proximal half medially offset 30� from its longitudinalaxis). These features, in combination with the presence of“clubbed” dorsal vertebrae (?pathological), a sacrum composedof eight vertebrae, whose last sacrals have a neural spine about2.5 times the height of the centrum, and a distal end of the ischiumthat forms a hypertrophied foot-like expansion, make it possible todistinguish Pararhabdodon from all other hadrosauroids.In order to test the relationships of Pararhabdodon isonensis,

it was included in the analysis made by Horner et al. (2004:supplement). Our analysis found 504 most parsimonious treesof 190 steps each (CI = 0.636; RI = 0.738). The strict consensustree shows a highly unresolved topology; the 50% majority-ruleconsensus tree supports the inclusion of Pararhabdodon withinHadrosauridae as a basal member of the clade (SupplementaryData 1, www.vertpaleo.org/jvp/JVPcontents.html). P. isonensisshares at least two hadrosaurid synapomorphies: elevation ofthe cervical zygapophyseal peduncles on neural arches to extendwell above the level of the neural canal, long and dorsally archedpostzygapophyses (character 74 of Horner et al., 2004), and adorsoventrally narrow proximal scapula with an acromion pro-cess that projects horizontally, and a reduced area for the articu-lation with the coracoid (character 80). Moreover, it apparentlyshares with the Lambeosaurinae-Hadrosaurinae clade the pres-*Corresponding author.

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ence of at least eight sacral vertebrae (character 75 of Horneret al., 2004). The number of sacral vertebrae is unknown inTelmatosaurus. Pararhabdodon has a low and gently roundedmaxillary apex in lateral view, a character interpreted as synapo-morphic for Lambeosaurinae by Horner et al. (2004: character48), but regarded as plesiomorphic by Prieto-Marquez et al.(2006: character 41) and Evans & Reisz (2007: character 20).Pararhabdodon lacks unambiguous characters of derived Lam-beosaurinae: i.e., posterior dorsal and sacral neural spines elon-gate to more than three times the height of the centrum(character 76 of Horner et al., 2004; ratio about 2.5 in IPSSRA 24).Inclusion of Pararhabdodon in recent phylogenetic analyses

does not give conclusive results. On the basis of the data fromGodefroit et al. (2008: appendix 3), Pararhabdodon is placed inpolytomy with Aralosaurus and Wulagasaurus as the sister groupto lambeosaurines and hadrosaurines (strict consensus of 36 treesof 81 steps each; CI = 0,913; RI = 0,962) (Supplementary Data 2,www.vertpaleo.org/jvp/JVPcontents.html). However, when in-cluded in the analysis of Evans & Reisz (2007: appendix 2),Pararhabdodon is placed together with Aralosaurus as the basal-most members of Lambeosaurinae (strict consensus of 10 trees of114 steps each; CI = 0,886; RI = 0,916) (Supplementary Data 3,

www.vertpaleo.org/jvp/JVPcontents.html). Pararhabdodon sha-res with lambeosaurines an expanded deltopectoral crest on thehumerus and a large and pendent foot on the ischium. However,these characters are also present on some non-hadrosauridmembers of Hadrosauroidea, and therefore they can haveindependently evolved in P. isonensis (Prieto-Marquez et al.,2006).In conclusion, Pararhabdodon is a poorly understood taxon

that has an unresolved position within Hadrosauridae.- Koutalisaurus kohlerorum, Les Llaus (Lleida). This taxon waserected on the basis of an edentulous right dentary (IPS SRA27) from Les Llaus, near Abella de la Conca (Tremp Basin,Lleida). This material comes from a similar geographic area andfrom the same unit that yielded the remains of P. isonensis,which is Early Maastrichtian to early Late Maastrichtian in age(Lopez-Martınez, 2003). The new taxon was diagnosed byPrieto-Marquez et al. (2006) on the basis of a single character:“edentulous portion of the dentary that is extremely elongatedand medially projected, so that the distance between the sym-physis and the lateral wall of the element is three times itsmediolateral breadth”.

Koutalisaurus kohlerorum exemplifies the problem of ere-cting new species on the basis of very fragmentary material.

FIGURE 1. A, Location map of Iberian hadrosauroid sites discussed in the text; B, MPZ 2005/90, Lambeosaurinae indet., right ilium from Blasi-3(Aren, Huesca) in lateral view; C, D, MPZ 99/664-668, Lambeosaurinae indet., cranial remains from Blasi-1 (Aren, Huesca): C, reconstructed skull inlateral view showing preserved elements: left jugal, fragment of maxilla, dentary and surangular (inversed); D, left dentary in medial view; E, F,MGUV 2200 and MPV 181, Hadrosauridae indet., right dentaries from La Solana (Tous, Valencia) in medial views; G, IPS 36338, Hadrosauroideaindet., left dentary from Fontllonga (Lleida) in medial view. Scales are equal to 5 cm. All the remains are Late Maastrichtian in age.

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Prieto-Marquez et al. (2006) chose the ‘splitting option’ becauseit is “as susceptible to error as lumping IPS SRA 27 and theremaining material into one taxon”. However, the validity ofK. kohlerorum is conditional upon the discovery of additionalmaterial.

Koutalisaurus kohlerorum was regarded by Prieto-Marquezet al. (2006) as a hadrosaurid of uncertain phylogenetic position.It shares with the Lambeosaurinae-Hadrosaurinae clade a toothrow parallel with the lateral side of the dentary in occlusal view(Prieto-Marquez et al., 2006: character 8.2, supplementary data;but note that this character may be ontogenetic; P. Godefroit,pers. comm.). Moreover, the diastema between the first dentarytooth and the predentary is extremely long, as in Edmontosaurusand closely related forms, but unlike hadrosaurines the numberof tooth positions in the dentary tooth row is less than 40(Godefroit et al., 2008: characters 38, 41). More recently,Prieto-Marquez et al. (2007) have reported the discovery of skullremains, a dentary, teeth and postcranial bones from the Maas-trichtian of Lleida that may represent K. kohlerorum. On thebasis of jugal and maxillary features, they provisionally referredthis material to the Lambeosaurinae.- Lambeosaurinae indet., Blasi-3 specimen, Aren (Huesca). Cru-zado-Caballero et al. (2005) described a complete right iliumfrom the Blasi-3 locality of Aren (lower part of the ConquesFormation, Tremp Group; upper Maastrichtian). MPZ 2005/90is a small-sized ilium (length: 46,8 cm), which possesses a mas-sive supracetabular process and a strongly curved preacetabularprocess (Fig. 1B). It shares with the Lambeosaurinae-Had-rosaurinae clade a sigmoidal dorsal margin that is distinctly de-pressed over the supracetabular process and dorsally bowed overthe base of the preacetabular process (Horner et al., 2004: char-acter 100; Prieto-Marquez et al., 2006: character 116, supplemen-tary data). The Blasi-3 ilium is deep, with the depth of theproximal region of the preacetabular process more than 50 percent the depth of the central body of the ilium, as is typicalin lambeosaurines except Lambeosaurus and Corythosaurus(Prieto-Marquez et al., 2006: character 118, supplementary da-ta). MPZ 2005/90 resembles the ilia of the lambeosaurinesCorythosaurus, Olorotitan and Parasaurolophus in having a pre-acetabular process that is twice as long as the postacetabularprocess (Prieto-Marquez et al., 2006: character 117, supplemen-tary data).- Lambeosaurinae indet., Blasi-1 specimens, Aren (Huesca).Cranial bones from the Blasi-1 site of Aren (MPZ 99/664-667)consist of a jugal, a fragmentary maxilla with teeth, a dentarywith the dental battery, and a surangular (Lopez-Martınez et al.,2001). All the bones but the surangular come from the left sideof the cranium, and probably belong to the same individual(Fig. 1C, D). The Blasi-1 material was provisionally referred toEuhadrosauria indet. on the basis of both jugal and dentaryfeatures (Lopez-Martınez et al., 2001). These authors consideredthat MPZ 99/664-667 has a mosaic of both hadrosaurine andlambeosaurine features. In fact, the jugal shows a combinationof characters only known in lambeosaurines: a truncated anddorsally expanded rostral process; V-shaped posteroventralflange, forming a pointed corner; very slight constriction or un-constricted quadratojugal process (Prieto-Marquez et al., 2006:characters 28, 30, supplementary data; see also Evans & Reisz,2007: characters 28, 31; Godefroit et al., 2008: character 30).Unlike derived hadrosaurids, the number of functional teethper tooth position is only two in the Blasi-1 dentary, but thismight be ontogenetic (two functional teeth are known in youngspecimens of Olorotitan versus three in adults; P. Godefroit,pers. comm.). The lateral surface of the dentary is parallel withthe symphysis, as in Koutalisaurus, and the number of toothpositions is roughly similar in both dentaries (33 in Blasi-3, 35in Koutalisaurus). In contrast to Koutalisaurus, the length of theedentulous portion of Blasi-1 (measured between the first den-

tary tooth and the predentary joint) does not reach half thelength of the dental battery. Moreover, MPZ 99/665 differs fromall other dentaries found in the Iberian Peninsula, includingthose of Tous and Fontllonga, in having only a single carina onthe teeth; secondary ridges are absent. The possibility that thespecimens from Blasi-1 and Blasi-3 belong to the same taxon isopen, but the remains cannot be directly compared.- Hadrosauridae indet., La Solana specimens, Tous (Valencia).Two well preserved dentaries (MGUV 2200, MPV 181) anda fragmentary one (MPV 182) have been found in the LaSolana site of Tous (Valencia), one of them described by Com-pany et al. (1998) (Figs. 1E, F). The fossiliferous beds, datedbiostratigraphically as Late Maastrichtian, can be correlatedwith the Villalba de la Sierra Formation of the SE Iberian Range(eastern Spain). The dentaries and in situ teeth are alike andpresumably belong to the same taxon. Differences in size andin the number of tooth positions are probably due to ontogen-etic changes. The Tous taxon shares with the Hadrosauridaethe presence of a tooth row that terminates posterior to theapex of the coronoid process (You et al., 2003: character 29;Horner et al., 2004: character 10; Prieto-Marquez et al., 2006:character 6). MPV 181 has 36-37 tooth positions in the dentalbattery (Fig. 1F), as in hadrosaurines and lambeosaurines, butunlike them it retains a subvertical coronoid process (Company,2004; see Godefroit et al., 1998: character 35; Prieto-Marquezet al., 2006: character 7). This plesiomorphic condition is alsoobserved in MGUV 2200 (Fig. 1E). The lanceolate teeth areornamented with one well-developed carina plus a faint sec-ondary ridge on the mesial region of the crown (Companyet al., 1998). On the basis of these features, the Tous taxon isconsidered to be a basal hadrosaurid that is more derived thanTelmatosaurus.- Hadrosauroidea indet., Fontllonga specimen, Fontllonga(Lleida). IPS 36338 (Fig. 1G) consists of a left dentary with partof the dental battery, but the symphyseal region is broken(Casanovas et al., 1999b). The fossil was recovered near thetown of Fontllonga in the uppermost Maastrichtian strata of theTremp Group, close to the presumed Cretaceous-Tertiaryboundary. Casanovas et al. (1999b) concluded that the Fontl-longa taxon is more derived than is Telmatosaurus but lacks anumber of features that diagnose the Hadrosaurinae-Lambeo-saurinae clade. Nevertheless, a reappraisal of the material indi-cates that it might belong to a derived non-hadrosauridhadrosauroid instead of a basal hadrosaurid. In IPS 36338, thetooth row terminates ventral to the apex of the coronoid processand not posterior to it, unlike Telmatosaurus and the hadrosaur-ine-lambeosaurine clade (Horner et al., 2004: character 10;Prieto-Marquez et al., 2006: character 6). Moreover, the Fontl-longa taxon and Telmatosaurus differ from typical euhadrosaur-ians in that they have a subvertical coronoid process, thetooth row is not parallel with the lateral side of the dentary,and the number of tooth positions in the dental battery isless than 34 (Prieto-Marquez et al., 2006: characters 7, 8, 11;Godefroit et al., 2008: characters 39, 41). In the Fontllongataxon, each tooth position consists of two functional teeth andtwo replacement teeth. The jaw displays an abnormal tooth re-placement pattern: the distal teeth are no more completelyerupted and worn than the central ones, and the rows subject tothe same replacement are not step-like but more or less horizon-tal; the significance of this feature is obscure (Ostrom, 1961).The teeth have a well-developed ridge plus a vestigial, faint sec-ondary one.

LATE MAASTRICHTIAN HADROSAUROIDASSEMBLAGES

Outside the Iberian Peninsula, Telmatosaurus transsylvanicusis known in the Lower Maastrichtian to the base of the Upper

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Maastrichtian in the Hateg and Transylvanian basins of Roma-nia (Weishampel et al., 1993; Dalla Vecchia, 2006). Other hadro-sauroid remains are known in the Maastrichtian rocks ofsouthern France, Germany, Belgium, The Netherlands, Slove-nia, and the Ukraine (see Laurent, 2003; Dalla Vecchia, 2006and references inside). The occurrence of Pararhabdodon insouthern France is under debate (Prieto-Marquez et al., 2006),but cannot be excluded (Laurent, 2003). With the exception of afew euhadrosaurian remains from the Dutch Limburg, includingteeth (Weishampel et al., 1999; Jagt et al., 2003), the remainderof the material has been referred to Hadrosauridae indet. Mostof these remains are in need of revision.The hadrosauroid assemblages from the Late Maastrichtian of

Europe are very different from those known in the other land-masses of the Northern Hemisphere. Both in North Americaand Asia, the associations comprise exclusively euhadrosaurians(see Table). In western North America (Canada: Alberta, Sas-katchewan; USA: Colorado, Montana, North and South Dakota,Texas and Wyoming), the assemblages are dominated by hadro-saurines (Edmontosaurus sp., ?Gryposaurus), whereas lambeo-saurines are very rare or absent (?Lambeosaurus in the LanceFm. of Wyoming and ?Parasaurolophus in the Hell Creek Fm. ofMontana are listed by Weishampel et al., 2004; but see Godefroitet al., 2001, 2008). In eastern Asia (eastern Russia, northeasternChina), the assemblages are dominated by lambeosaurines(Amurosaurus, Charonosaurus, Olorotitan, Sahaliyania), buthadrosaurines (Kerberosaurus, Wulagasaurus) are also repre-sented (Godefroit et al., 2001, 2003, 2004, 2008; Bolotskyand Godefroit, 2004). By contrast, the Late Maastrichtianassemblages of Iberia consist of both derived euhadrosaurians(e.g., the lambeosaurines of Blasi-1 and Blasi-3) and basalhadrosaurids (the Tous taxon). The possible occurrence of non-hadrosaurid Hadrosauroidea (Fontllonga taxon) is worthy ofconsideration.

PALEOBIOGEOGRAPHICAL IMPLICATIONS

The diversity and affinities of the Iberian hadrosauroidsshould be taken into account when reconstructing the paleobio-geography of the European dinosaurs. The primitive characterof the hadrosaurid faunas of Europe has been explained in termsof geographical isolation (as evolutionary relicts) during theLate Cretaceous (Weishampel et al., 1993; Casanovas et al.,1999b). At this time, hadrosaurids had a large geographical dis-tribution: Asia, Europe, western and eastern North America,South America and Antarctica (Horner et al., 2004; Weishampelet al., 2004). Fossil evidence supports a Laurasian origin forhadrosaurids (and hadrosauroids), but a North American originhas also been proposed for the Hadrosauridae, with dispersalevents to Asia, Europe and South America (Horner et al.,2004). On the basis of a tentative reconstruction of ancestral

areas, Prieto-Marquez et al. (2006) considered the origin of theHadrosauridae to be equally parsimonious in North America,Asia and Europe.An Asian origin for the hadrosauroids is a reasonably hypoth-

esis (see also You et al., 2003; Godefroit et al., 2008). Averianovand Archibald (2005) have suggested that primitive hadrosaurids(together with basal zhelestid mammals) may have reached Eur-ope from Asia before the formation of the Turgai Strait in theTuronian. Subsequent isolation of the European archipelagofrom Asia and other landmasses may have facilitated the surviv-al of basal hadrosaurids (e.g., Telmatosaurus) in the Maastrich-tian of Europe. In fact, the relationships of Telmatosaurus andEuhadrosauria imply high minimal divergence time values (ap-proximately 20 Myr; see Weishampel et al., 1993). However,Martin et al. (2005) have hypothesized a high-latitude dispersal(Thulean route) for latest Cretaceous terrestrial vertebratesfrom eastern North America to Europe on the basis of the oc-currence of herpetotheriid marsupials in the type section of theMaastrichtian of The Netherlands. Sea level and climatic condi-tions might have been sufficiently favorable to allow such aNorth Atlantic dispersal route, e.g. immigration of hadrosauridsfrom North America to Europe (Martin et al., 2005; contraHead, 1998). If so, derived hadrosaurids, such as lambeosaur-ines, could have reached the European archipelago during thelatest Cretaceous.

CONCLUSIONS

A review of the hadrosauroid record from the latest Creta-ceous of Spain indicates a greater diversity than previouslythought, with at least three Late Maastrichtian taxa, including aLambeosaurinae member from the Blasi site, in Aren (Huesca),a basal hadrosaurid from La Solana, in Tous (Valencia), and apossible non-hadrosaurid member of the Hadrosauroidea fromFontllonga (Lleida). The only genera and species described todate are Pararhabdodon isonensis Casanovas et al., 1993 andKoutalisaurus kohlerorum Prieto-Marquez et al., 2006, bothfrom the Lower Maastrichtian to the lower Upper Maastrichtianof the Tremp-Isona area in Lleida. The membership of Para-rhabdodon in Lambeosaurinae has been questioned, but its rela-tionships within Hadrosauridae are not yet fully resolved. Thephylogenetic position of Koutalisaurus (a possible junior syno-nym of Pararhabdodon) is uncertain within Hadrosauridae.The hadrosauroid assemblage from the Upper Maastrichtian

of the Iberian Peninsula is very different from those known inNorth America and Asia, which are dominated by euhadrosaur-ians: lambeosaurines in eastern Asia and hadrosaurines in west-ern North America. By contrast, the Iberian faunas (and those ofEurope as a whole) consist of both derived and basal forms. In abiogeographical context, the primitive character of some Euro-pean hadrosauroids (e.g., Telmatosaurus transsylvanicus and the

TABLE. Late Maastrichtian hadrosauroid taxa from the Northern Hemisphere.

Western North America Europe Asia

Edmontosaurus regalis (H) Lambeosaurinae indet. (Blasi-1, Blasi-3) Amurosaurus riabinini (L)Edmontosaurus annectens* (H) Hadrosauridae indet. (Tous) Charonosaurus jiayinensis (L)Edmontosaurus saskatchewanensis (H) ?Hadrosauroidea indet. (Fontllonga) Kerberosaurus manakini (H)Edmontosaurus sp. (H) Pararhabdodon isonensis*** (St. Roma) Olorotitan arhanensis (L)?Gryposaurus sp. (H) Koutalisaurus kohlerorum*** (Les Llaus) Sahaliyania elunchunorum (L)?Lambeosaurus sp.** (L) Wulagasaurus dongi (H)?Parasaurolophus sp.** (L)

Data from Horner et al. (2004), Godefroit et al. (2004, 2008); Weishampel et al. (2004); Lund & Gates (2006), and this work. In round brackets,Iberian sites. H, hadrosaurine; L, lambeosaurine.*Including Anatotitan copei.**Data from Weishampel et al. (2004). The presence of Lambeosaurinae in Late Maastricthian sites of North America is doubtful (Godefroit et al.,2001, 2008).***The age of these taxa is uncertain: Early Maastrichtian to early Late Maastrichtian (Lopez-Martınez, 2003).

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Fontllonga taxon) could be explained in terms of geographicalisolation during the Late Cretaceous, and the occurrence ofderived euhadrosaurians (e.g., lambeosaurines) by a late disper-sion from Asia and/or North America.

ACKNOWLEDGMENTS

XPS acknowledges the financial support provided by theSpanish Ministerio de Educacion y Ciencia (MEC) (ProgramaRamon y Cajal; projects CGL2004-02338 and CGL2007-64061/BTE), the UPV/EHU (9/UPV 00121.310-15303/2003) and theBasque Government (CIC07/14-361). Research work of JIC andPCC was supported by the MEC and the ERDF (CGL 2004-03393, CGL2007-62469/BTE) and the Government of Aragon(Grupos Consolidados 2007, Department for Education andCulture). Research of JIR-O is supported by the Regional Min-istry for Culture and Tourism of the Principality of Asturias(Protocolo CN-04-226). The manuscript was significantly im-proved thanks to the helpful reviews from P. Godefroit andF. DallaVecchia. Z. Herrera and I. Perez-Urresti (Univ. Zara-goza) prepared some of the photographs. A. Galobart (IPS,Sabadell) and A. Prieto-Marquez (Florida State University,Tallahassee) facilitated information to us.

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Submitted July 21, 2008; accepted October 25, 2008.

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