New Material and Phylogenetic Position of Arenysaurus ardevoli, a Lambeosaurine Dinosaur from the Late Maastrichtian of Arén (Northern Spain)

Abstract

Arenysaurus ardevoli is a lambeosaurine hadrosaurid from the late Maastrichtian of Ar´en (Huesca, northern Spain) that has recently been described. The holotype is the first and the most complete lambeosaurine with a braincase from Europe. In this paper, we present a complete description of the postcranial skeleton, which was poorly described when the taxon was named because it was partially unprepared, and new information on several cranial bones (jugal, maxilla, and dentition). A new phylogenetic analysis of Arenysaurus and the closely related Blasisaurus canudoi, also from the late Maastrichtian of Ar´ en, places them inside the Parasaurolophini in a dichotomy with Parasaurolophus spp. Paleobiogeographically, the presence of Arenysaurus and its relationships with other lambeosaurines suggest at least one geodispersal event from Asia to Europe no later than the middle–late Campanian.

Full text

Journal of Vertebrate Paleontology 33(6):1367–1384, November 2013
©2013 by the Society of Vertebrate Paleontology
ARTICLE
NEW MATERIAL AND PHYLOGENETIC POSITION OF ARENYSAURUS ARDEVOLI,
A LAMBEOSAURINE DINOSAUR FROM THE LATE MAASTRICHTIAN OF AR ´
EN
(NORTHERN SPAIN)
PEN ´
ELOPE CRUZADO-CABALLERO,*,1,†JOS ´
E IGNACIO CANUDO,1MIGUEL MORENO-AZANZA,1
and JOS ´
E IGNACIO RUIZ-OME ˜
NACA1,2
1Grupo Aragosaurus-IUCA, ´
Area de Paleontolog´
ıa, Facultad de Ciencias, Universidad de Zaragoza, Pedro Cerbuna 12, 50009
Zaragoza, Spain, http://www.aragosaurus.com; penelope@unizar.es; jicanudo@unizar.es; mmazanza@unizar.es;
2Museo del Jur´
asico de Asturias (MUJA), 33328 Colunga, Spain, jigruiz@unizar.es
ABSTRACT—Arenysaurus ardevoli is a lambeosaurine hadrosaurid from the late Maastrichtian of Ar ´
en (Huesca, northern
Spain) that has recently been described. The holotype is the first and the most complete lambeosaurine with a braincase from
Europe. In this paper, we present a complete description of the postcranial skeleton, which was poorly described when the taxon
was named because it was partially unprepared, and new information on several cranial bones (jugal, maxilla, and dentition).
A new phylogenetic analysis of Arenysaurus and the closely related Blasisaurus canudoi, also from the late Maastrichtian of
Ar´
en, places them inside the Parasaurolophini in a dichotomy with Parasaurolophus spp. Paleobiogeographically, the presence
of Arenysaurus and its relationships with other lambeosaurines suggest at least one geodispersal event from Asia to Europe no
later than the middle–late Campanian.
SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP
INTRODUCTION
Hadrosaurids were common herbivorous dinosaurs during the
Late Cretaceous in Laurasia (Horner et al., 2004). Their distri-
bution was nearly worldwide, but the best-known fossil record
comes from North America and Asia (Lund and Gates, 2006).
In the last 15 years, numerous new remains have been disco-
vered in Europe, which, together with the basal hadrosaurid Tel-
matosaurus Nopcsa, 1903,show the rich paleobiodiversity of the
Campanian–Maastrichtian interval (Dalla Vecchia, 2009a, 2009b;
Pereda-Suberbiola et al., 2009; Prieto-M´
arquez and Wagner, 2009;
Cruzado-Caballero et al., 2010a, 2010b).
In Europe, the most abundant fossil record of hadrosaurids is
from the Maastrichtian of Spain. Remains from the Tremp Basin
(south-central Pyrenees) have made possible the description of
three lambeosaurine taxa (Pararhabdodon isonensis Casanovas-
Cladellas, Santaf´
e-Llopis, and Isidro-Llorens, 1993 [including
Koutalisaurus kohlerorum Prieto-M´
arquez, Gaete, Rivas, Galo-
bart, and Boada, 2006]; Arenysaurus ardevoli Pereda-Suberbiola,
Canudo, Cruzado-Caballero, Barco, L ´
opez-Mart´
ınez, Oms,
and Ruiz-Ome ˜
naca, 2009; and Blasisaurus canudoi Cruzado-
Caballero, Pereda-Suberbiola, and Ruiz-Ome ˜
naca, 2010a) and
an indeterminate hadrosaurine (Pereda-Suberbiola et al., 2009;
Prieto-M´
arquez and Wagner, 2009; Cruzado-Caballero et al.,
2010a, 2010b). The European hadrosaurid record thus provides
proof of high and previously unknown vertebrate paleodiversity in
this continent during the latest Cretaceous. This fauna has affini-
ties with North American and Asian taxa (Pereda-Suberbiola,
2009) as a result of several geodispersal events that occurred in
the course of the latest Cretaceous. Here we present a complete
*Corresponding author. †Current address: CONICET-Instituto de In-
vestigaci ´
on en Paleobiolog´
ıayGeolog
´
ıa, R´
ıo Negro, Argentina.
description of several new cranial remains and the postcranial
skeleton of Arenysaurus ardevoli from the late Maastrichtian site
of Blasi 3. In a previous paper, Pereda-Suberbiola et al. (2009)
erected the taxon, described part of the cranial material, and
reported briefly on the postcranial remains. New remains, which
were unprepared in 2009, are here described for the first time
(i.e., jugal, caudal vertebrae, ribs, hemal arches, and pubis). Thus,
Arenysaurus (Fig. 1) is the best-represented and most complete
lambeosaurine from the Ibero-Armorican Island. This taxon
fills a gap in our understanding of the phylogeny of European
lambeosaurines and has provided new data about geodispersal
events with the rest of Laurasia.
Institutional Abbreviations—MDE,Mus´
ee des Dinosaures, Es-
peraza, France; MPZ, Museo Paleontol ´
ogico de la Universidad de
Zaragoza, Zaragoza, Spain.
GEOLOGIC AND CHRONOLOGIC FRAMEWORK
The Late Cretaceous vertebrate-bearing localities of Blasi are
located 2 km to the west of the town of Ar´
en (or Areny de
Noguera, Huesca Province) in northeastern Spain. Several pub-
lications have given an account of the vertebrate assemblages at
these localities (L ´
opez-Mart´
ınez et al., 2001; Pereda-Suberbiola
et al., 2009; Blain et al., 2010; Cruzado-Caballero et al., 2010a,
2010b).
Seven of the localities have produced hadrosaurid remains
(Blasi 1, Blasi 2a, Blasi 2b, Blasi 3, Blasi 3.4, Blasi 4, and Blasi
5). These are located stratigraphically in the Ar´
en Formation
(Blasi 1), the La Posa Formation (Blasi 2a, Blasi 2b, Blasi 3,
and Blasi 3.4), and the Conques Formation (Blasi 4 and Blasi 5).
In terms of the regional geology, they are located in the Tremp
Basin of the south-central Pyrenees (Riera et al., 2009), or the
Tremp-Graus Basin (Pujalte and Schmidt, 2005). It should be
1367
Downloaded by [University of Zaragoza], [Miguel Moreno-Azanza] at 08:09 12 November 2013

1368 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 33, NO. 6, 2013
FIGURE 1. Arenysaurus ardevoli. Com-
posite cranial reconstruction in left lat-
eral view, with the following elements:
MPZ2008/1, skull; MPZ2008/256, left maxilla;
MPZ2011/01, left jugal; MPZ2008/258, left
dentary; MPZ2008/259, right surangular
(reversed).
noted that in previous papers the sites Blasi 2a to Blasi 5 were
placed in the Tremp Formation following L ´
opez-Mart´
ınez et al.
(2001). Nevertheless, according to Pujalte and Schmidt (2005:79),
“the term Tremp Group reflects more appropriately than the for-
mer Tremp Formation the stratigraphic complexity of the so-
called <Garumnian>of the south Pyrenean Tremp-Graus basin.”
The age of the ‘Garumnian’ in the south-central Pyrenees ranges
from Maastrichtian (and probably latest Campanian; see Lopez-
Mart´
ınez et al., 2001:55) to lower Eocene (Claret Formation in
Pujalte and Schmidt [2005]; ‘fluvial upper red unit’ of the Tremp
Formation in Riera et al. [2009]).
In accordance with the lithostratigraphic diagram of the Tremp
Group including the Ar´
en/Areny section (Pujalte and Schmidt,
2005:fig. 1B; see also Riera et al., 2009:fig. 3), in this paper we place
Blasi 2a, Blasi 2b, Blasi 3, and Blasi 3.4 in the lower gray part of the
Tremp Group, in the La Posa Formation (‘marine-to-continental
transitional gray unit’ of the Tremp Formation, sensu Riera et al.,
2009), and Blasi 4 and Blasi 5 in the lower red part of the Tremp
Group, in the Conques Formation (‘fluvial lower red unit’ of the
Tremp Formation, sensu Riera et al., 2009).
Using magnetostratigraphy, the Blasi sites have been placed in
the upper part of a normal polarity chron correlated with chron
C30n (Oms and Canudo, 2004; Pereda-Suberbiola et al., 2009:fig.
2), i.e., they are younger than 67.7 Ma and slightly older than 65.8
Ma (Ogg et al., 2008).
Besides hadrosaurids, remains of other vertebrates have
been found at these sites: fishes, amphibians, lizards, turtles,
crocodylians, and theropod and sauropod dinosaurs (Table 1). Re-
cently, Pu´
ertolas et al. (2011) described a new crocodylian named
Arenysuchus gascabadiolorum. This crocodylian comes from a
new site (the El´
ıas site in the Conques Formation) discovered in
2008, very close to the sites of Blasi 1–3.
The Blasi 3 site, from which the Arenysaurus material was re-
covered, has provided remains of at least two other hadrosaurid
individuals, an indeterminate juvenile lambeosaurine and a
small adult hadrosaurid (Cruzado-Caballero and Canudo, 2005;
Cruzado-Caballero et al., 2005; Cruzado-Caballero, 2012).
SYSTEMATIC PALEONTOLOGY
DINOSAURIA Owen, 1842
ORNITHISCHIA Seeley, 1887
ORNITHOPODA Marsh, 1881
HADROSAURIDAE Cope, 1870
LAMBEOSAURINAE Parks, 1923
ARENYSAURUS Pereda-Suberbiola, Canudo,
Cruzado-Caballero, Barco, L ´
opez-Mart´
ınez, Oms, and
Ruiz-Ome ˜
naca, 2009
ARENYSAURUS ARDEVOLI Pereda-Suberbiola, Canudo,
Cruzado-Caballero, Barco, L ´
opez-Mart´
ınez, Oms, and
Ruiz-Ome ˜
naca, 2009
(Figs. 1–11)
Original Diagnosis—Lambeosaurine hadrosaurid characterized
by a very prominent frontal dome, more developed than in
other adult specimens; nearly vertical prequadratic process of the
squamosal and jugal process of the postorbital; deltopectoral crest
of the humerus oriented anteriorly. It differs from other lam-
beosaurines in having a unique combination of characters: short
frontal, with a posterior length/maximal width ratio estimated at
0.5; midline ridge of parietal approximately at the level of the
Downloaded by [University of Zaragoza], [Miguel Moreno-Azanza] at 08:09 12 November 2013

CRUZADO-CABALLERO ET AL.—ARENYSAURUS ARDEVOLI FROM SPAIN 1369
TABLE 1. Faunal list from the Blasi sites.
Blasi site
Fauna 1 2 (2a and 2b) 3 3.4 4 5
Actinopterygii
Lepisosteidae indet. X
Pycnodontiformes indet. X
Teleostei indet. X
Amphibia
Albanerpetontidae
Albanerpeton aff. nexuosum X
Anura
Discoglossidae aff. Paradiscoglossus sp. X
Palaeobatrachidae indet. X
Squamata
Indeterminate lizard 1 (Iguania or
Scleroglossa)
X
Indeterminate lizard 2 (Scleroglossa,
Scincomorpha)
X
Anguidae indet. X
Alethinophidia indet. X
Chelonii
Solemydidae sp. X
Bothremydidae indet. X X X X
Crocodyliformes
Crocodyliformes indet. X X X
‘Trematochampsidae’ indet. X X
Eusuchia indet. X
cf. Arenysuchus gascabadiolorum X
Alligatoroidea indet. X X
Acynodon sp. X
Dinosauria
Theropoda
Theropoda indet. X X X
Coelurosauria indet. X
cf. Euronychodon sp. X
cf. Dromaeosauridae indet. X X
Maniraptora indet. X
Ornithopoda
Hadrosauridae indet. X X X X X
Euhadrosauria indet. X X X
Hadrosaurinae indet. X
Lambeosaurinae indet. XX
Lambeosaurinae
Arenysaurus ardevoli X
Blasisaurus canudoi X
Modified from L ´
opez-Mart´
ınez et al. (2001), with data from Torices et al. (2004), Murelaga and Canudo (2005), Pereda-Suberbiola et al. (2009), Blain
et al. (2010), Cruzado-Caballero et al. (2010a, 2010b, 2013), and Pu´
ertolas et al. (2011).
postorbital-squamosal bar; parietal not interposed between the
squamosals in the occipital surface of the skull; lateral side of
squamosal relatively low above the cotyloid cavity.
Holotype—MPZ2008/1, a partial, articulated skull comprising
the skull roof and braincase.
Paratypes—Cranial material: MPZ2008/256, fragmentary left
maxilla; MPZ2008/257, fragmentary right maxilla; MPZ2008/258,
left dentary with 12 teeth; MPZ2008/259, right surangular;
MPZ2008/260–263, four isolated teeth. Postcranial skele-
ton: MPZ2007/706, MPZ2007/954–955, MPZ2008/264–267,
seven cervical vertebrae; MPZ2008/268, a dorsal vertebra;
MPZ2008/269–270, two dorsal ribs; MPZ2008/271, partial
sacrum with ossified tendons; MPZ2004/480, pathological cau-
dal vertebra; MPZ2006/20, 14 articulated caudal vertebrae
and hemal arches; MPZ2008/272, MPZ2008/313, two caudal
vertebrae; MPZ2008/314, MPZ2008/330, two hemal arches;
MPZ2008/331–332, two ossified tendons; MPZ2008/333a–333b,
right scapula (two fragments); MPZ2008/334, right coracoid;
MPZ2008/336, right humerus; MPZ2008/335, fragmentary right
ilium; MPZ2007/707, right pubis; MPZ2007/711, right femur;
MPZ2008/337, left femur.
Referred Specimens—MPZ2011/1, left jugal; MPZ2012/767,
cervical vertebrae; MPZ2012/746–747, MPZ2012/767, three cer-
vical ribs; MPZ2012/748–753, six dorsal ribs; MPZ2008/275–276,
MPZ2008/281, MPZ2008/286, MPZ2008/288, MPZ2008/290–292,
MPZ2008/296, MPZ2008/298–300, MPZ2008/302, MPZ2008/304,
MPZ2008/306, MPZ2008/309–312, MPZ2012/754–758, 24 caudal
vertebra; MPZ2008/315–317, MPZ2008/319, MPZ2008/321–323,
MPZ2008/325–326, MPZ2008/328–329, 11 hemal arches.
Comment—Given their association and size, it is assumed that
all the paratypes from the original publication (Pereda-Suberbiola
et al., 2009) and the newly referred specimens (topotypes) come
from a single hadrosaurid skeleton, from which the holotype
derives. However, because all the material is partially disar-
ticulated and comes from a locality that preserves two other
small hadrosaurid individuals (i.e., an indeterminate juvenile lam-
beosaurine and a small adult hadrosaurid; Cruzado-Caballero and
Canudo, 2005; Cruzado-Caballero et al., 2005, Cruzado-Caballero,
Downloaded by [University of Zaragoza], [Miguel Moreno-Azanza] at 08:09 12 November 2013

1370 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 33, NO. 6, 2013
2012), it is preferable to designate a holotype and not convert all
the material (paratypes and topotypes) assigned to Arenysaurus
into a single holotype.
DESCRIPTION
For the skull and dentary descriptions, see Pereda-Suberbiola
et al. (2009:546–567, figs. 2–3). Other cranial remains are described
and figured below for the first time.
New Cranial Material
Jugal—MPZ2011/1 is a fragmentary left jugal (Fig. 2A, B).
Its morphology is similar to those of Blasisaurus and an inde-
terminate lambeosaurine from Blasi 4 (MPZ2008/1884; Cruzado-
Caballero et al., 2010b). The jugal includes the preserved dorsal
half of the anterior process, which is expanded dorsally, unlike
the elongated and slender anterior processes of Telmatosaurus
and Tethyshadros Dalla Vecchia, 2009b (Weishampel et al.,
1993; Dalla Vecchia, 2009b). The posterodorsal side of the an-
terior process is less inclined anteriorly than that in Blasisaurus,
MPZ2008/1884, and MPZ2008/1885 (Hadrosaurine indet. from
the Blasi 5 site; Cruzado-Caballero et al., 2010b). This form of the
anterior process is similar to Velafrons Gates, Sampson, Delgado
de Jes´
us, Zanno, Eberth, Hernandez-Rivera, Aguill ´
on Mart´
ınez,
and Kirkland, 2007, and Lambeosaurus Parks, 1923 (Evans and
Reisz, 2007; Gates et al., 2007). A medially and slightly anteri-
orly projecting maxillary process is present on the anterior pro-
cess of the jugal, unlike Blasisaurus. The postorbital process is
triangular in cross-section, robust, and oriented posterodorsally,
forming an angle of 58◦with the long axis of the jugal, similar to
those in Blasisaurus,Parasaurolophus sp., and Velafrons, with an
angle of 60◦; and unlike Amurosaurus Godefroit, Bolotsky, and
Van Itterbeeck, 2004, with an angle of 45◦;Corythosaurus Brown,
1914, with an angle of 75◦;andLambeosaurus and MPZ2008/1885,
with an angle of 80◦(Ostrom, 1961; Godefroit et al., 2004; Evans
and Reisz, 2007; Evans et al., 2007; Gates et al., 2007; Cruzado-
Caballero et al., 2010a, 2010b). The orbital fenestra is ‘V’-shaped,
unlike Blasisaurus and MPZ2008/1884 (Cruzado-Caballero et al.,
2010a, 2010b).
Maxilla—Two maxillary fragments (MPZ2008/256-257, left and
right respectively; Fig. 3A–D) have been found in close association
with the skull and are here referred to Arenysaurus. MPZ2008/256
is a left maxillary fragment. It is very long anteroposteriorly
(227 mm) and narrow lateromedially. The premaxillary shelf is
wide, concave, and rough. Posterior to this shelf, on the dorsal sur-
face, there is a maxillary foramen. In medial view, there are almost
15 special foramina interconnected by a gently curving horizontal
groove, along the entire length of the bone as preserved.
MPZ2008/257 is a right maxillary fragment with an anteropos-
terior length of 176 mm. The maxilla is broken medially, and
the tooth positions can be observed. There are 14 tooth po-
sitions preserved, and these are slightly concave anteriorly. In
medial view, there are 14 special foramina interconnected by a
gently curving horizontal groove. The ectopterygoid ridge seems
very prominent on the lateral side of the maxilla. This ectoptery-
goid ridge turns ventrally, unlike in Pararhabdodon,inwhichit
is horizontal (Prieto-M´
arquez et al., 2006). The maxillary fora-
men is present close to the posterior border of the premaxillary
shelf.
Surangular—MPZ2008/259 is a right surangular (Fig. 4A–D).
It is anteroposteriorly longer and dorsoventrally wider than that
of Blasisaurus (Cruzado-Caballero et al., 2010a). The surangular
lacks a foramen on the lateral body of the bone, as is typical
in hadrosaurids. In dorsal view, it is laterally very convex. The
surangular portion of the mandibular glenoid is cup-shaped and is
shallow and expanded both anteroposteriorly and mediolaterally.
A deep groove for the M. pterygoideus dorsalis is located under
the glenoid and near a prominent ridge that separates it from
the angular facet (Ostrom, 1961). The lateral lip is robust. The
retroarticular process is elongate and strong, it is compressed
FIGURE 2. Arenysaurus ardevoli left jugal (MPZ2011/1) in medial (A)andlateral(B)views.Abbreviations:ap, anterior process; lp, lacrimal process;
mp, maxillary process; pp, postorbital process.
Downloaded by [University of Zaragoza], [Miguel Moreno-Azanza] at 08:09 12 November 2013

CRUZADO-CABALLERO ET AL.—ARENYSAURUS ARDEVOLI FROM SPAIN 1371
FIGURE 3. Arenysaurus ardevoli maxillae in medial (A, C)andlateral(B, D)views.A,B, right maxilla (MPZ2008/257); C,D. left maxilla
(MPZ2008/256). Abbreviations:ectr, ectopterygoid ridge; pmxs, premaxillary shelf; sf, special foramina.
Downloaded by [University of Zaragoza], [Miguel Moreno-Azanza] at 08:09 12 November 2013

1372 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 33, NO. 6, 2013
FIGURE 4. Arenysaurus ardevoli right surangular (MPZ2008/259) in medial (A), lateral (B), dorsal (C), and ventral (D)views.Abbreviations:af,
angular facet; mg, mandibular glenoid; rp, retroarticular process.
mediolaterally, and on the ventral side there is an insertion area
for the M. pterygoideus ventralis (Ostrom, 1961).
Isolated Teeth—The maxillary teeth are narrower than the den-
tary tooth. One isolated right dentary tooth (MPZ2008/263), two
right maxillary teeth (MPZ2008/260, MPZ2008/262), and one frag-
mentary tooth (MPZ2008/261) have been collected (Fig. 5A–C).
The crowns have the typical lanceolate shape of hadrosaurids
(Horner et al., 2004). In both types of teeth there is a strong and
straight median primary ridge on the enameled side of the crown,
and the mesial and distal edges are not denticulate, as in Bla-
sisaurus and unlike Telmatosaurus and Tethyshadros (Weisham-
pel et al., 1993; Dalla Vecchia, 2009b; Cruzado-Caballero et al.,
2010a). The height/width ratio is about 3.2 in the dentary tooth
and 3.1 in the maxillary teeth. The dentary and maxillary teeth
possess a faint secondary ridge mesial to the primary ridge.
Axial Skeleton
Pereda-Suberbiola et al. (2009:567) provided only short descrip-
tions of the cervical, dorsal, sacral, and caudal vertebrae, and none
of them were figured. Below, the vertebral series is described and
figured for the first time in its entirety.
Cervical Vertebrae and Ribs—Eight partial to complete cervi-
cal vertebrae have been found (MPZ2007/706, MPZ2007/954–955,
MPZ2008/264–267, MPZ2012/767; Fig. 6A–C, Table 2). These
vertebral elements are indistinguishable from those of all other
hadrosaurids. The centra are strongly opisthocoelous and partic-
ularly wide and short, with the following proportions: width >
height =length, and with the heart-shaped anterior and posterior
ends typical of the hadrosaurids (Horner et al., 2004). The ante-
rior articular surface is globular, and the posterior surface is wider
and cup-shaped. On the ventral side there is a prominent longi-
tudinal keel. Few nutritional foramina are present on the lateral
surfaces of the centra. The parapophyses are located at midheight
on the depressed lateral surfaces. These are short and are linked to
the cervical ribs. The cervical ribs are dorsoventrally narrow and
curved backwards. The neural arch surrounds a large neural canal.
The lateral transverse processes are not very long, and are robust
and curved backwards with a triangular cross-section. Near to the
base of the transverse process there is a wide prezygapophysis with
a flat surface facing upwards and inwards. The postzygapophy-
seal processes are long, have a triangular cross-section, and are
curved backwards and outwards. They extend well above the level
of the neural canal. The width between the postzygapophyseal
processes decreases posteriorly in the cervical series. On the top
of the postzygapophyseal process in ventral view there is a large
circular postzygapophysis with flat surface face.
Three of the cervical vertebrae (MPZ2007/954, MPZ2007/955,
MPZ2008/265) lack the neural spine. MPZ2007/706 has a neural
spine that is curved backwards, mediolaterally compressed, an-
teroposteriorly wide, and dorsoventrally short, and that does not
extend beyond the postzygapophyseal processes.
Several fragmentary cervical ribs have been recovered with the
typical morphology of hadrosaurids (Horner et al., 2004). They
are dorsoventrally flat and curved backwards. The tuberculum is
expanded dorsoventrally at the joint with the cervical vertebrae.
It is shorter than the capitulum, and MPZ2012/767 preserves an
anteriorly projecting spine.
Dorsal Vertebrae and Ribs—A single dorsal vertebra has been
found (MPZ2008/268; Fig. 6D–F, Table 3). As with the cervical
vertebrae, this vertebra is indistinguishable from those of all other
hadrosaurids. This vertebra is slightly deformed obliquely and
eroded, so nutritional foramina cannot be observed. The trans-
verse processes are long, have a cross-section that is triangular
at the base and ellipsoidal at the end, and curve backwards and
outwards. Near to the base of the transverse process there is a
wide prezygapophysis with flat surface face. The location of the
prezygapophysis indicates an anterior position in the dorsal ver-
tebra series. In the neural spine, the postzygapophyseal processes
are near the base in ventral view. They are long, ellipsoidal, and
Downloaded by [University of Zaragoza], [Miguel Moreno-Azanza] at 08:09 12 November 2013

CRUZADO-CABALLERO ET AL.—ARENYSAURUS ARDEVOLI FROM SPAIN 1373
FIGURE 5. Arenysaurus ardevoli teeth. A, right dentary tooth (MPZ2008/263) in lingual view; B, right maxillary tooth (MPZ2008/260) in labial view;
C, detail of the secondary ridge in labial view. Abbreviation:sr, secondary ridge. A,B, sacle bar equals 1 cm. C, scale bar equals 0.5 cm.
have a flat surface that faces upwards and inwards. Between the
postzygapophyseal processes there is a deep groove.
The neural spine is elongated, mediolaterally compressed, acute
dorsally, and approximately 3.5 times the height of the centrum.
It is inclined backwards, with the anterior border convex and the
posterior slightly concave in lateral view.
The dorsal ribs (MPZ2008/269–270, MPZ2012/748–753) are
fragmentary and, like the cervical ribs, have the typical
hadrosaurid morphology (Horner et al., 2004). The lengths of the
shafts are variable, and the cross-section is dorsoventrally flat.
Sacral Vertebrae—MPZ2008/271 is a partial sacrum formed
from five fused centra, whose dorsal part is completely destroyed.
An isolated element is identified as the last sacral vertebra
(Fig. 6G, H). The articular surfaces are elliptical in outline, wider
than high, and concave. In ventral view, a broad and shallow sul-
cus is present in the sacrum, as in Amurosaurus,Bactrosaurus
Gilmore, 1933, and Pararhabdodon (Godefroit et al., 1998, 2004;
Casanovas-Cladellas et al., 1999). In Arenysaurus, the groove is
not present in the first and last vertebrae. The neural spines are
broken, except for the neural spine of the last centrum, which has
a height that is more than three times that of the centrum, as in
lambeosaurines (Horner et al., 2004).
Caudal Vertebrae—About two-thirds of the caudal series is
preserved. Fifty-four vertebrae, including 14 that are articulated
(MPZ2006/20), have been found, all of them indistinguishable
from those of all other hadrosaurids. These vertebrae represent
TABLE 2. Measurements of cervical vertebrae, in mm.
Specimen A-P length D-V height Lt-M width
Angle between postzygapophyseal
processes
Length transverse
processes (R/L)
MPZ2007/706 75.54 58.46 65.81 105◦74.5/73.85
MPZ2007/955 58.39∗49.39 78.97 90◦46.81∗/43.88∗
MPZ2008/267 X X X 90◦21.74∗/28.57∗
MPZ2008/266 X X X X 27.94∗/X
MPZ2007/954 68.41∗43.26∗50.72∗85◦31.67∗/39.85∗
MPZ2008/264 54.87∗55.41 55.46 X X
MPZ2008/265 80.04∗48.33∗47.51 65◦39.88/16.92∗
MPZ2012/767 X X X 91◦50/X
Abbreviations:A-P, anteroposterior; D-V, dorsoventral; L, left; Lt-M, lateromedial; R,right;∗, estimated; X, not preserved.
Downloaded by [University of Zaragoza], [Miguel Moreno-Azanza] at 08:09 12 November 2013

1374 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 33, NO. 6, 2013
FIGURE 6. Arenysaurus ardevoli vertebrae in anterior (A,D,G), posterior (B), lateral (Cand F), and ventral (Gand E)views.A–C, cervical
vertebrae (MPZ2007/955); E–G, dorsal vertebrae (MPZ2008/268); G,H, sacral vertebrae 1–5 (MPZ2008/271). Abbreviation:s, sulcus.
Downloaded by [University of Zaragoza], [Miguel Moreno-Azanza] at 08:09 12 November 2013

CRUZADO-CABALLERO ET AL.—ARENYSAURUS ARDEVOLI FROM SPAIN 1375
TABLE 3. Measurements of dorsal vertebrae, in mm.
Specimen A-P length D-V height Lt-M width Length spine Angle spine
Length transverse
processes (R/L)
Angle transverse
processes (R/L)
MPZ2008/268 69.13∗40.43∗41.46 172∗71◦95.99/67.83∗84◦/X
Abbreviations:A-P, anteroposterior; D-V, dorsoventral; L, left; Lt-M, lateromedial; R,right;∗, estimated; X, not preserved. Angle spine, angle between
the neural spine and the vertical axis; Angle transverse processes, angle between the transverse process and the sagittal plane.
almost all of the anterior and middle sections of the tail. The
anterior-most centra are typically amphiplatyan, anteroposteriorly
narrow, with hexagonal articular surfaces, a transverse process,
and hemapophyseal facets in the ventral side. Posteriorly, they
present a sequence of dorsoventral reduction and anteroposte-
rior increase in the dimensions of the centra; the mid-posterior
vertebrae lack hemapophyseal facets. In lateral view, the sides of
the mid-posterior centra are slightly depressed, lack transverse
processes, and are pierced by irregularly distributed nutritional
foramina of variable size.
The neural spines are very long (more than three times the
height of the centrum), robust, and subvertical (with an angle to
the vertical axis of almost 90◦) in the anterior-most vertebrae. In
the posterior-most vertebrae, the neural spines are slender, steeply
inclined backwards (angle between 34◦and 45◦), and their end is
slightly curved dorsally. The tip of the neural spine in the anterior-
most vertebrae is broken, and we do not know if these were
slightly expanded transversally as in Barsboldia Maryanska and
Osm ´
olska, 1981, and Amurosaurus (Maryanska and Osm ´
olska,
1981; Godefroit et al., 2004).
In the middle caudal series, five vertebrae in the block
MPZ2006/20 show marks parallel to the posterodorsal axis that
are probably due to predation. These marks coincide in position
with the pathological vertebra MPZ2004/480 (Canudo et al., 2005).
The neural spine of MPZ2004/480 presents a swelling and devia-
tion near to the tip that produces a distinctive convexity on the
right side and concavity on the left side (Fig. 7A–D). In anterior
view, the anomalous bone growth creates an oval hole in the cen-
tral section. The left side exhibits at least two parallel fractures
partially covered by anomalous bone growth. These fractures have
an anteroposterior orientation, and one is located at the base of
the anomalous bone development, whereas the second is located
above the hole. Canudo et al. (2005) explained these pathologies
as being the result of a probable attack by a theropod dinosaur and
the subsequent infection of the injury.
Hemal Arches—The hemal arches have the typical hadrosaurid
‘Y’ shape, are long and mediolaterally flat, and the articular facets
are separated by an open hemal canal (Horner et al., 2004). The
angle between the hemal arches and the centra cannot be mea-
sured because the hemal arches were discovered disarticulated
from the caudal series.
Appendicular Skeleton
Pereda-Suberbiola et al. (2009) and Cruzado-Caballero et al.
(2009) provided short descriptions of the right scapula, humerus,
and pubis, and the femora. Only one of these bones, the right
humerus, was figured (Pereda-Suberbiola et al., 2009:fig. 5). All
FIGURE 7. Arenysaurus ardevoli pathology caudal vertebra (MPZ204/480) in anterior (A), posterior (B), left lateral (C), and right lateral (D)views.
Abbreviations:pat, pathology.
Downloaded by [University of Zaragoza], [Miguel Moreno-Azanza] at 08:09 12 November 2013

1376 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 33, NO. 6, 2013
FIGURE 8. Arenysaurus ardevoli right scapula in medial (Aand B) and dorsal (C)views.A, proximal fragment (MPZ2008/333a); B,C, distal fragment
(MPZ2008/333b). Abbreviations:cf, coracoid facet; dr, deltoid ridge; g, glenoid.
of these bones and new unpublished appendicular remains are de-
scribed and figured below.
Scapula—MPZ2008/333a and MPZ2008/333b are two frag-
ments of a broken right scapula: the proximal fragment and an
almost complete blade, respectively (estimated length 525 mm;
Fig. 8A–C). MPZ2008/333a is narrow and has a rectangular shape,
unlike the triangular shape of Telmatosaurus (Weishampel et al.,
1993). The deltoid fossa is broad and bears numerous longitudinal
striations and a prominent knob, probably an extensive attach-
ment site for a powerful M. supracoracoideus (Dilkes, 2000). This
knob and the anterior orientation of the deltopectoral crest would
have increased the power of the animal’s arm. The acromion pro-
cess is horizontally projecting, similar to Wulagasaurus Godefroit,
Hai, Yu, and Lauters, 2008, and Secernosaurus Brett-Surman,
1979 (Godefroit et al., 2008; Prieto-M´
arquez and Salinas, 2010). It
is less robust than in Pararhabdodon (Prieto-Marquez et al., 2006)
and parallel to the dorsal side, unlike in Tethyshadros (Dalla
Vecchia, 2009b). The acromion process bears striations for the
attachment of the M. trapezius (Godefroit et al., 2004). The cora-
coid facet is large, rough, and cup-shaped, as in Charonosaurus
Godefroit, Zan, and Jin, 2000, and Amurosaurus (Godefroit et al.,
2001, 2004). The glenoid facet is smaller, straight, and oriented
backwards. These two facets form an angle of approximately 135◦,
similar to the condition in Wulagasaurus and unlike MDE-Les-25
(Euhadrosauria indet. from Lestaillats, France), which has an
angle of 90◦,andPararhabdodon, which has an angle of 100◦
(Laurent et al., 1999; Prieto-M´
arquez et al., 2006; Godefroit et al.,
2008).
MPZ2005/333b possesses a narrow scapular neck, as in lam-
beosaurines and in contrast to more primitive hadrosaurids
(Horner et al., 2004), so that the blade is approximately 60%
wider than the proximal neck. The scapular blade is narrow and
in lateral view it is anteriorly convex. The dorsal and ventral mar-
gins of the scapula are curved slightly downward and diverge,
with the maximum width on the symmetrical posterior side, as
in Amurosaurus,Lambeosaurus,Nipponosaurus Nagao, 1936, Sa-
haliyania Godefroit, Hai, Yu, and Lauters, 2008, and an indetermi-
nate lambeosaurine from New Mexico (Nagao, 1936; Williamson,
2000; Godefroit et al., 2004, 2008; Suzuki et al., 2004; Evans and
Reisz, 2007). According to Godefroit et al. (2001, 2004), the elon-
gation and ventral curvature of the scapular blade may be related
to a general increase in the power of the forelimb, because this
lengthens the in-lever arms of the M. teres major, which inserted
along the posteroventral side of the scapular blade.
Coracoid—MPZ2008/334 is a right coracoid (Fig. 9). As is
common in hadrosaurids, the hook-shaped ventral process is
prominent and points anteroventrally (Horner et al., 2004;
Godefroit et al., 2008). The coracoid foramen is large and
elliptical; it is completely surrounded by the coracoid. It is an-
teroposteriorly wider than dorsoventrally high, but less so than
in Charonosaurus (Godefroit et al., 2001). The anterior edge
of the coracoid is thinner than the posterior one. The ventral
edge forms a small glenoid fossa. On the posterior side, the
scapular articular surface and the glenoid have a similar length,
but the scapular articulation is thicker than the glenoid. This
surface is thicker dorsoventrally than mediolaterally, unlike in
Hadrosaurus Leidy, 1858, Brachylophosaurus Sternberg, 1953,
Edmontosaurus,Saurolophus Brown, 1912, and Gryposaurus
Lambe, 1914 (Prieto-M´
arquez et al., 2006). The two articular
surfaces form an angle of about 135◦, this angle being greater than
those of the Pararhabdodon (100◦), Brachylophosaurus (115◦),
Bactrosaurus and Amurosaurus (120◦), and Secernosaurus (126◦),
and less than those of Charonosaurus and Nipponosaurus (150◦)
(Godefroit et al., 1998, 2001, 2004; Suzuki et al., 2004; Prieto-
M´
arquez et al., 2006; Prieto-M´
arquez, 2007; Prieto-M´
arquez and
Salinas, 2010). The biceps tubercle is large and projects laterally.
Humerus—MPZ2008/336 is an incomplete right humerus (see
Pereda-Suberbiola et al., 2009:fig. 5). It is massive, the cross-
section is rectangular, and the shaft is straight, unlike in Tel-
matosaurus (Weishampel et al., 1993) and Tethyshadros (Dalla
Downloaded by [University of Zaragoza], [Miguel Moreno-Azanza] at 08:09 12 November 2013

CRUZADO-CABALLERO ET AL.—ARENYSAURUS ARDEVOLI FROM SPAIN 1377
FIGURE 9. Arenysaurus ardevoli right cora-
coid (MPZ2008/334) in anterior view. Abbre-
viations:bt; biceps tubercle; cf, coracoid fora-
men; gf, glenoid fossa; scf, scapular facet; vp,
ventral process.
Vecchia, 2009b). The deltopectoral crest is oriented anteriorly as
in Wulagasaurus, so that the maximum width of the shaft is dis-
tal unlike in other lambeosaurines, where the maximum width is
at the level of the deltopectoral crest (Godefroit et al., 2008). The
bicipital groove is deep and narrow. The proximal portion of the
humerus is broken, and the distal portion presents the ulnar and
radial condyles. The ulnar condyle is more developed than the ra-
dial one. The intercondylar groove is slightly wider on the poste-
rior side than on the anterior side. On the medial side, there is a
prominent knob on the shaft.
Ilium—MPZ2008/335 is the preacetabular process of a
right ilium (Fig. 10A, B). It is wide dorsoventrally, narrow
mediolaterally, and ventrally directed. In medial view, near to the
dorsal side there is a groove between two ridges, and a slight scar
for the M. iliotibialis (Dilkes, 2000).
Pubis—MPZ2007/707 is the prepubic blade of a robust right
pubis (Fig. 11A, B). It is anteroposteriorly short, dorsoventrally
expanded, and is nearly symmetrical. It is similar to the type 5
pubis (Parasaurolophus Parks, 1922, and Bactrosaurus)ofBrett-
Surman and Wagner (2006) and to stage 1 (Parasaurolophus)
of Prieto-M´
arquez (2010). In lateral view, it is slightly concave
and has strong muscular marks that posteriorly are perpendicu-
lar to the neck and anteriorly are oblique to the blade. In me-
dial view, there are also strong muscular marks, probably for
the M. ambiens (Brett-Surman and Wagner, 2006). The neck
is wide and strong, with the dorsal side nearly straight. The
Downloaded by [University of Zaragoza], [Miguel Moreno-Azanza] at 08:09 12 November 2013

1378 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 33, NO. 6, 2013
FIGURE 10. Arenysaurus ardevoli right ilium (MPZ2008/335) in lateral
(A) and medial (B)views.
iliac peduncle is strong, anteriorly directed, and has a triangular
cross-section.
Femora—Both femora (MPZ2007/711 and MPZ2008/337) were
recovered from the site. MPZ2007/711 is a complete right femur
with a typical hadrosaurid form and a length of 711 mm. It has
a straight massive shaft, unlike Telmatosaurus, which has a dis-
tal portion that is medially inclined (Weishampel et al., 1993).
The cross-section is rectangular and mediolaterally narrow. The
femoral head and greater trochanter are separated by a wide
groove. The greater trochanter is narrow anteroposteriorly and
wider laterally than the femoral head. A groove separates the
greater trochanter from the lesser trochanter, which is developed
mediolaterally. Below the lesser trochanter there is a slight inser-
tion mark for the M. puboischiofemoralis internus 2 (pars dor-
salis) (Dilkes, 2000). The fourth trochanter is triangular in shape
and mediolaterally narrow. In lateral view, the insertion area for
the M. caudifemoralis brevis (Dilkes, 2000) is not separated into
two areas, unlike the femora from Blasi 1 (Cruzado-Caballero
et al., 2009). The distal condyles are expanded anteroposteriorly
and proximodistally. They are lateromedially narrow due to diage-
netic deformation; the intercondylar extensor groove is not closed,
probably due to this deformation. MPZ2008/337 is an incomplete
left femur. Only the distal part has been preserved, and its dia-
physis is partially hollow. The distal condyles are bioturbated by
invertebrate galleries, as is the case in MPZ2007/711. They are
wide proximodistally and they are not complete anteroposteriorly.
There is a well-developed intercondylar extensor groove that is
closed, forming a tunnel (Horner et al., 2004).
Bioturbation has been observed in both femora and also in two
other bones (MPZ2008/336, right humerus; MPZ2008/333a, right
scapula). These are possibly traces produced by necrophagous in-
sects (Cruzado-Caballero et al., in preparation).
PHYLOGENETIC ANALYSIS
In this paper, we follow the definition of Sereno (1998:62)
for the clade Hadrosaurinae, which is “all hadrosaurids closer
to Saurolophus than to Parasaurolophus,” and for Lambeosauri-
nae, which is “all hadrosaurids closer to Parasaurolophus than
to Saurolophus.” Within Lambeosaurinae there are two widely
recognized lineages, which were defined by Evans and Reisz
(2007:385) as Parasaurolophini: “taxa more closely related to
Parasaurolophus walkeri than to Corythosaurus casuarius”; and
Corythosaurini: “taxa more closely related to C.casuarius than to
P.walkeri.”
To assess the phylogenetic relationships of the Spanish lam-
beosaurines, we have included these taxa in the matrix proposed
by Godefroit et al. (2012), to which we have added one more
character from Prieto-M´
arquez (2010, character 82), which scores
important information for differentiating Arenysaurus and Bla-
sisaurus. In this data set, we have corrected some typos related to
the Spanish hadrosaurids (see Appendix 1) and included new ma-
terial described in the present work. Character distributions were
analyzed with Mesquite 2.74 (Maddison and Maddison, 2010). The
resulting matrix included 22 taxa coded for 119 characters and
was run with TNT version 1.1 (Goloboff et al., 2008). All char-
acters were unordered and equally weighted. A heuristic search
with 1000 replications, retaining 10 trees per replication, was car-
ried out using Probactrosaurus Rozhdestvensky, 1966, as the out-
group. Five most parsimonious trees were obtained (tree length =
188 steps, consistency index =0.713, retention index =0.805,
rescaled consistency index =0.574). Bremer supports and boot-
strap values were calculated for each branch to assess its robust-
ness.
The strict consensus tree is presented in Figure 12. In contrast
to Godefroit et al. (2012), we have not deleted Sahaliyania a pos-
teriori, because its position remained stable in all trees, forming
a trichotomy with the tribes Parasaurolophini and Lambeosaurini
in the consensus. We also considered the deletion of Velafrons,
because its variable position within Lambeosaurini has been re-
ported in previous studies (Godefroit et al., 2012). If Velafrons is
removed, only one tree of 186 steps is recovered. Nevertheless, we
have chosen not to remove it, because its position is not relevant
for the current study. The resulting topology resembles previously
published hypotheses, with the exceptions of the polytomy within
Lambeosaurini attributable to Velafrons and, more significantly,
the position of Arenysaurus and Blasisaurus, which has turned out
to be more derived than previously stated.
Previous analyses reported Arenysaurus as the sister taxon
to Amurosaurus and all more derived lambeosaurines (Pereda-
Suberbiola et al., 2009; Godefroit et al., 2012). Cruzado-Caballero
et al. (2010a) found the clade formed by Arenysaurus and Bla-
sisaurus to be placed in a trichotomy with Amurosaurus and the
node that includes Lambeosaurini and Parasaurolophini. With the
inclusion of the new postcranial data we have recovered a more
derived position for the Arenysaurus +Blasisaurus clade, nested
within the tribe Parasaurolophini. Although the three synapomor-
phies of Parasaurolophini cannot be coded for either Arenysaurus
or Blasisaurus,Arenysaurus shares the following characters with
the genus Parasaurolophus: (1) a caudal process of the postorbital
elongated above the infratemporal fenestra (character 36:1); (2)
a short diastema between the first dentary tooth and the preden-
tary, also shared with Blasisaurus (character 58:0); (3) mediolat-
erally broad distal condyles of the humerus (character 78:1); and
(4) a deltoid ridge of the scapula that is dorsoventrally deep and
craniocaudally long, with a well-demarcated ventral margin (char-
acter 109:1). Nevertheless, both taxa are well differentiated from
all other Parasaurolophini in having a wide lingual projection of
the symphyseal region of the dentary (character 105:0). It is of
Downloaded by [University of Zaragoza], [Miguel Moreno-Azanza] at 08:09 12 November 2013

CRUZADO-CABALLERO ET AL.—ARENYSAURUS ARDEVOLI FROM SPAIN 1379
FIGURE 11. Arenysaurus ardevoli right pubis (MPZ2007/707) in medial (A)andlateral(B)views.Abbreviations:ilp, iliac peduncle; preb, prepubic
blade.
interest to compare Arenysaurus and Blasisaurus with Pararhab-
dodon, the only other Spanish lambeosaurine. Arenysaurus differs
in that it has (1) sacral neural spines elongated, approximately
three times the height of the centrum or greater (character 68:1);
(2) humeral distal condyles mediolaterally broad, which flare mod-
erately from the shaft of the humerus (character 78:1); (3) a lingual
projection of the symphyseal region of the dentary, with a ratio
between the labiolingual extension of the symphyseal region and
the maximum labiolingual width of the dentary that reaches 1.65
(character 105:0); and (4) a concave medial or lateral profile of the
dorsal margin of the rostral edentulous region of the dentary for
articulation with the predentary (character 106:0). Of these char-
acters, Blasisaurus shares with Arenysaurus its differences from
Pararhabdodon in the characters 58, 105, and 106.
Blasisaurus also shares with all other derived lambeosaurids a
straight dentary tooth row (character 119:1), a derived state not
Downloaded by [University of Zaragoza], [Miguel Moreno-Azanza] at 08:09 12 November 2013

1380 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 33, NO. 6, 2013
FIGURE 12. Strict consensus of five most
parsimonious trees (tree length =188 steps,
consistency index =0.713, retention index =
0.805, rescaled consistency index =0.574).
Data set based on the matrix of Godefroit et al.
(2012), with the addition of character 48 from
Prieto-Marquez (2010), and the correction of
several typos with respect to the Spanish taxa.
See Appendix 1 and Supplementary Materi-
als for character scores and data matrix. Note
the position of Arenysaurus and Blasisaurus
as members of Lambeosaurini, instead of the
more basal position recovered in previous
studies, as sister taxa of Amurosaurus (Pereda
Suberbiola et al., 2009; Cruzado-Caballero
et al., 2010a; Godefroit et al., 2012). Num-
bers over the branches are Bremer support val-
ues and numbers under the branches represent
bootstrap values after 1000 replicates.
shared with Arenysaurus, which possesses the ancestral condition
of a dentary tooth row that is bowed lingually.
PALEOBIOGEOGRAPHIC IMPLICATIONS
The geodispersal events of hadrosaurids between western North
America and eastern Asia, in both directions, have been amply
proved in previous works (Head, 1998, 2001; Godefroit et al., 2003,
2004, 2008; Horner et al., 2004; Fiorillo, 2008; Sues and Averianov,
2009; Prieto-M ´
arquez, 2010). Similarly, a connection between Asia
and Europe during the second half of the Late Cretaceous has
been discussed (Pereda-Suberbiola et al., 2009; Prieto-M´
arquez
and Wagner, 2009; Cruzado-Caballero et al., 2010b).
According to Pereda-Suberbiola et al. (2009) and Prieto-
M´
arquez and Wagner (2009), among other authors, the connec-
tion between Asia and Europe was probably interrupted prior to
the Maastrichtian. Before that, there were ‘semipermeable’ barri-
ers as a result of land bridges that emerged or were submerged de-
pending on climatic and sea-level changes. These bridges allowed
several dinosaur taxa to use the European archipelago as a refuge;
among these were the lambeosaurines (Pereda-Suberbiola, 2009).
Conversely, several tetrapod taxa from the Late Cretaceous
of Europe are considered to have Euramerican affinities: pa-
leobatrachid frogs, batrachosauroidid salamanders, solemydid
turtles, basal crocodyloids, and, tentatively, chelydroid turtles,
alligatoroid crocodyliforms, and nodosaurid dinosaurs (see refer-
ences in Pereda-Suberbiola, 2009; Pu´
ertolas et al., 2011). These
taxa probably used bridges that sporadically opened high-latitude
routes from eastern North America into Europe and allowed
faunal exchange between the two areas (see Pu´
ertolas et al., 2011,
and references therein). Our phylogenetic hypotheses suggest
that these geodispersal events also affected the European fauna
of lambeosaurine hadrosaurids.
The distribution of lambeosaurines in a simplified phylogenetic
tree shows three hypothetical geodispersal episodes (see Fig. 13):
(1) Pararhabdodon or its ancestors migrated from Asia to the
Iberian island of the European archipelago (Prieto-M´
arquez and
Wagner, 2009); (2) Arenysaurus,Blasisaurus, or their ancestors
migrated from Asia to the Iberian island; and (3) Parasaurolo-
phus spp. or their ancestors migrated probably from Asia to North
America.
Geodispersal events 1 and 2 (from Asia to Europe) could
have occurred at the same time, no later than the middle–late
Downloaded by [University of Zaragoza], [Miguel Moreno-Azanza] at 08:09 12 November 2013

CRUZADO-CABALLERO ET AL.—ARENYSAURUS ARDEVOLI FROM SPAIN 1381
FIGURE 13. Biogeographic implications of the lambeosaurine phylogenetic analysis carried out in this paper. Geodispersal events are marked with
numbers: 1, 2, from Asia to Europe; 3 from Asia to North America. Dashed-line boxes represent uncertain ranges of presence of species. Boundaries
(Ma) after Walker and Geissman (2009), and the age of the represented species is based on Prieto-M´
arquez (2010).
Downloaded by [University of Zaragoza], [Miguel Moreno-Azanza] at 08:09 12 November 2013

1382 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 33, NO. 6, 2013
Campanian according to Prieto-M´
arquez and Wagner (2009).
Geodispersal event 3 (from Asia to North America) probably oc-
curred prior to or during the early–middle Campanian.
Two additional geodispersal events have been recognized in a
previous paper by the present authors (Cruzado-Caballero et al.,
2011:fig. 2), based on the phylogenetic analyses of Evans and Reisz
(2007) and Gates et al. (2007) with the addition of Arenysaurus
and Blasisaurus:(4)Velafrons and Lambeosaurus spp. or their an-
cestors migrated from Asia to North America before or during
the early Campanian; and (5) Olorotitan or its ancestors migrated
from North America to Asia in the middle–late Campanian. Nev-
ertheless, the phylogeny of corythosaurines is not resolved in the
present phylogenetic analysis, and these two geodispersal cannot
be tested.
Hypothetical geodispersal episodes 3–5 are for the moment ten-
tative. It is thus necessary to reach a greater consensus on the phy-
logeny of hadrosaurids and a greater knowledge of the European
taxa to test them.
CONCLUSIONS
Arenysaurus ardevoli is the most complete lambeosaurine
from the Iberian Peninsula, indeed from anywhere in Europe.
It possesses several cranial characters that differentiate it from
Pararhabdodon: jugal with anterior process expanded dorsally,
straight posterodorsal end, postorbital process forming an angle
of 58◦with the long axis of the jugal, and ‘V’-shaped orbital fen-
estra; maxilla with ectopterygoid ridge ventrally turned; dentary
with anterior portion modestly deflected ventrally, a moderate
diastema, and a lingual projection of the symphyseal region of
the dentary with a ratio between the labiolingual extension of
the symphyseal region and the maximum labiolingual width of the
dentary that reaches 1.65; the presence of a mesial secondary ridge
in maxillary and dentary teeth (Pereda-Suberbiola et al., 2009).
Additional postcranial characters that differentiate Arenysaurus
from Pararhabdodon are rectangular scapula with glenoid and
coracoid facets forming an angle of 135◦; straight humerus with
deltopectoral crest oriented anteriorly; sacral neural spines elon-
gated, approximately three times the height of the centrum or
greater; humeral distal condyles mediolaterally broad and flar-
ing moderately from the shaft of the humerus; and pubis of
type 5 (Parasaurolophus and Bactrosaurus) sensu Brett-Surman
and Wagner (2006) and stage 1 (Parasaurolophus) sensu Prieto-
M´
arquez (2010).
Arenysaurus and Blasisaurus form a clade that is placed inside
the Parasaurolophini tribe in a dichotomy with Parasaurolophus
spp. The presence of Arenysaurus,Blasisaurus,andPararhab-
dodon in Europe and Parasaurolophus spp. in North America sug-
gests the occurrence of at least two geodispersal events of Asian
lambeosaurines from Asia to Europe and North America. These
geodispersal events probably involved the use of ‘semipermeable’
barriers that served as temporary connections between Asia and
Europe from the middle–late Campanian. Moreover, the Euro-
pean fauna of lambeosaurine hadrosaurids could have been af-
fected by geodispersal events of other tetrapod faunas from east-
ern North America.
ACKNOWLEDGMENTS
Financial support was provided by the Spanish Ministerio de
Econom´
ıa y Competitividad (projects CGL2010-16447/BTE),
the Gobierno de Arag ´
on (Grupos Consolidados, Departamento
de Educaci ´
on y Cultura; J.I.C., P.C.-C.), and Principado de
Asturias (University of Oviedo protocol CN-04-226; J.I.R.-O.).
The excavations and restorations of the fossils were subsidized by
the Gobierno de Arag ´
on, Diputaci´
on Provincial de Huesca, and
Ayuntamiento de Ar´
en. We acknowledge T. Gates (Ohio Univer-
sity College of Osteopathic Medicine, Athens, Ohio, and North
Carolina State University, Raleigh, North Carolina, U.S.A.) and
an anonymous reviewer for their comments on the manuscript.
The photographs were taken by Z. Herrera (University of
Zaragoza). R. Glasgow revised the English grammar.
LITERATURE CITED
Blain, H. A., J. I. Canudo, G. Cuenca-Besc ´
os, and N. L ´
opez-Mart´
ınez.
2010. Amphibians and squamates from the latest Maastrichtian (Up-
per Cretaceous) of Blasi 2 (Huesca, Spain). Cretaceous Research
31:433–446.
Brett-Surman, M. K. 1979. Phylogeny and paleobiogeography of
hadrosaurian dinosaurs. Nature 277:560–562.
Brett-Surman, M. K., and J. R. Wagner. 2006. Discussion of character anal-
ysis of the appendicular anatomy in Campanian and Maastrichtian
North American hadrosaurids—variation and ontogeny; pp. 125–169
in K. Carpenter (ed.), Horns and Beaks: Ceratopsian and Ornithopod
Dinosaurs. Indiana University Press, Bloomington and Indianapolis,
Indiana.
Brown, B. 1912. A crested dinosaur from the Edmonton Cretaceous. Bul-
letin of the American Museum of Natural History 31:131–136.
Brown, B. 1914. Corythosaurus casuarius, a new crested dinosaur from the
Belly River Cretaceous, with provisional classification of the family
Trachodontidae. Bulletin of the American Museum of Natural His-
tory 33:559–565.
Canudo, J. I., P. Cruzado-Caballero, and M. Moreno-Azanza. 2005. Pos-
sible theropod predation evidence in hadrosaurid dinosaurs from the
upper Maastrichtian (Upper Cretaceous) of Ar´
en (Huesca, Spain).
Kaupia. Darmst¨
adter Beitr¨
age zur Naturgeschichte 14:9–13.
Casanovas-Cladellas, M. L., J. V. Santaf´
e-Llopis, and A. Isidro-Llorens.
1993. Pararhabdodon isonense n. gen. n. sp. (Dinosauria). Estudio
morfol ´
ogico, radio-tomogr´
afico y consideraciones biomec´
anicas. Pa-
leontogia i Evoluci ´
o 26–27:121–131.
Casanovas-Cladellas, M. L., X. Pereda-Suberbiola, J. V. Santaf´
e, and
D.B. Weishampel. 1999. First lambeosaurine hadrosaurid from
Europe: palaeobiogeographical implications. Geological Magazine
136:205–211.
Cope, E. D. 1870. Synopsis of the extinct Batrachia, Reptilia, and Aves of
North America. Transactions of the American Philosophical Society
14:1–252.
Cruzado-Caballero, P. 2012. Restos directos de dinosaurios hadros ´
auridos
(Ornithopoda, Hadrosauridae) del Maastrichtiense superior
(Cret´
acico Superior) de Ar´
en (Huesca). Ph. D. thesis, Univer-
sity of Zaragoza, Zaragoza, Spain, 411 pp.
Cruzado-Caballero, P. and J. I. Canudo. 2005. Sobre la diversidad de los
hadrosaurios en el Maastricthiense superior (Cret´
acico superior) de
Ar´
en (Huesca); pp. 115–116 in E. Bern´
aldez, E. Mayoral and A.
Guerreiro dos Santos (eds.), XXI Jornadas de la Sociedad Espa ˜
nola
de Paleontolog´
ıa. Gesti ´
on e Investigaci ´
on de la Paleontolog´
ıa en el
Siglo XXI, Program and Abstracts, October 4–8, 2005, Sevilla.
Cruzado-Caballero, P., J. I. Canudo, and J. I.Ruiz-Ome ˜
naca. 2005. Nuevas
evidencias de la presencia de hadrosaurios lambeosaurinos (Di-
nosauria) en el Maastrichtiense superior de la Pen´
ınsula Ib´
erica
(Ar´
en, Huesca). Geogaceta 38:47–50.
Cruzado-Caballero, P., J. I. Canudo, and J. I. Ruiz-Ome ˜
naca. 2009. Los
f´
emures de Blasi (Ar´
en, Huesca, Spain): una contribuci ´
on a los
hadrosauroideos europeos del Maastrichtiense superior; pp. 197–205
in P. Huerta Hurtado and F. Torcida Fern´
andez-Baldor (eds.), Ac-
tas de las IV Jornadas Internacionales sobre Paleontolog´
ıa de Di-
nosaurios y su Entorno. Colectivo Arqueol ´
ogico y Paleontol ´
ogico de
Salas, Salas de los Infantes, Burgos, Spain.
Cruzado-Caballero, P., X. Pereda-Suberbiola, and J. I. Ruiz-Ome ˜
naca.
2010a. Blasisaurus canudoi gen. et sp. nov., a new lambeosaurine di-
nosaur (Hadrosauridae) from the latest Cretaceous of Ar´
en (Huesca,
Spain). Canadian Journal of Earth Sciences 47:1507–1517.
Cruzado-Caballero, P., J. I. Ruiz-Ome ˜
naca, and J. I. Canudo. 2010b.
Evidencias de la coexistencia de dinosaurios hadrosaurinos y lam-
beosaurinos en el Maastrichtiano superior de la Pen´
ınsula Ib´
erica
(Ar´
en, Huesca, Espa ˜
na). Ameghiniana 47:153–164.
Cruzado-Caballero, P., J. I. Canudo, M. Moreno-Azanza, and J. I. Ruiz-
Ome ˜
naca. 2011. The complex fauna of European Maastrichtian
Downloaded by [University of Zaragoza], [Miguel Moreno-Azanza] at 08:09 12 November 2013

CRUZADO-CABALLERO ET AL.—ARENYSAURUS ARDEVOLI FROM SPAIN 1383
hadrosaurids: contributions of the lambeosaurines from the Iberian
Peninsula; pp. 33–37 in D. R. Braman, D. A. Eberth, D. C. Evans, and
W. Taylor (eds.), International Hadrosaur Symposium at the Royal
Tyrrell Museum of Palaeontology, September 22–23, 2011. Program
and Abstracts. Royal Tyrrell Museum of Palaeontology, Drumheller.
Cruzado-Caballero, P., J. I. Ruiz-Ome ˜
naca, R. Gaete, V. Riera, O. Oms,
and J. I. Canudo. 2013. A new hadrosaurid dentary from the latest
Maastrichtian of the Pyrenees (north Spain) and the high diversity of
the duck-billed dinosaurs of the Ibero-Armorican Realm at the very
end of the Cretaceous. Historical Biology.
Dalla Vecchia, F. M. 2009a. European hadrosauroids; pp. 45–74 in P.
Huerta Hurtado and F. Torcida Fern´
andez-Baldor (eds.), Actas de
las IV Jornadas Internacionales sobre Paleontolog´
ıa de Dinosaurios y
su Entorno. Colectivo Arqueol ´
ogico y Paleontol ´
ogico de Salas, Salas
de los Infantes, Burgos, Spain.
Dalla Vecchia, F. M. 2009b. Tethyshadros insularis, a new hadrosauroid
dinosaur (Ornithischia) from the Upper Cretaceous of Italy. Journal
of Vertebrate Paleontology 29:1100–1116.
Dilkes, D. W. 2000. Appendicular myology of the hadrosaurian dinosaur
Maiasaura peeblesorum from the Late Cretaceous (Campanian) of
Montana. Transactions of the Royal Society of Edinburgh: Earth Sci-
ences 90:87–125.
Evans, D. C., and R. R. Reisz. 2007. Anatomy and relationships of Lam-
beosaurus magnicristatus, a crested hadrosaurid dinosaur (Ornithis-
chia) from the Dinosaur Park Formation, Alberta. Journal of Verte-
brate Paleontology 27:373–393.
Evans, D. C., R. R. Reisz, and K. Dupuis. 2007. A juvenile Parasaurolo-
phus (Ornithischia: Hadrosauridae) braincase from Dinosaur Provin-
cial Park, Alberta, with comments on crest ontogeny in the genus.
Journal of Vertebrate Paleontology 27:642–650.
Fiorillo, A. R. 2008. Dinosaurs of Alaska: implications for the Cretaceous
origin of Beringia; pp. 313–326 in R. B. Blodgett and G. D. Stanley Jr.
(eds.), The Terrane Puzzle: New Perspectives on Paleontology and
Stratigraphy from the North American Cordillera. Special Paper of
the Geological Society of America 442.
Gates, T. A., S. D. Sampson, C. R. Delgado de Jes ´
us, L. E. Zanno,
D. Eberth, R. Hern´
andez-Rivera, M. C. Aguill ´
on Mart´
ınez, and
J. I. Kirkland. 2007. Velafrons coahuilensis, a new lambeosaurine
hadrosaurid (Dinosauria: Ornithopoda) from the late Campanian
Cerro del pueblo Formation, Coahuila, M´
exico. Journal of Verte-
brate Paleontology 27:917–930.
Gilmore, C. W. 1933. On the dinosaurian fauna of the Iren Dabasu Forma-
tion. Bulletin of the American Museum of Natural History 67:23–78.
Godefroit, P., Y. Bolotsky, and V. Alifanov. 2003. A remarkable hollow-
crested hadrosaur from Russia: an Asian origin for lambeosaurines.
Comptes Rendus Palevol 2:143–151.
Godefroit, P., Y. L. Bolotsky, and I. Y. Bolotsky. 2012. Osteology and
relationships of Olorotitan arharensis, a hollow-crested hadrosaurid
dinosaur from the latest Cretaceous of Far Eastern Russia. Acta
Palaeontologica Polonica 57:527–560.
Godefroit, P., Y. L. Bolotsky, and J. Van Itterbeeck. 2004. The lam-
beosaurine dinosaur Amurosaurus riabinini, from the Maastrichtian
of Far Eastern Russia. Acta Palaeontologica Polonica 49:585–
618.
Godefroit, P., S. Zan, and L. Jin. 2000. Charonosaurus jiayinensis n. g., n.
sp., a lambeosaurine dinosaur from the late Maastrichtian of north-
eastern China. Comptes Rendus de l’Acad´
emie des Sciences. S´
erie
IIa, Sciences de la Terre et des plan`
etes 330:875–882.
Godefroit, P., S. Zan, and L. Jin. 2001. The Maastrichtian (Late Cre-
taceous) lambeosaurine dinosaur Charonosaurus jiayinensis from
north-eastern China. Bulletin de l’Institut Royal des Sciences Na-
turelles de Belgique, Sciences de la Terre 71:119–168.
Godefroit, P., S. Hai, T. Yu, and P. Lauters. 2008. New hadrosaurid di-
nosaurs from the uppermost Cretaceous of northeastern China. Acta
Palaeontologica Polonica 53:47–74.
Godefroit, P., Z.-M. Dong, P. Bultynck, H. Li, and L. Feng. 1998. Sino-
Belgian Cooperation Program “Cretaceous dinosaurs and mam-
mals from Inner Mongolia”; 1. New Bactrosaurus (Dinosauria,
Hadrosauroidea) material from Iren Dabasu (Inner Mongolia, P. R.
China). Bulletin de l’Institut Royal des Sciences Naturelles de Bel-
gique. Sciences de la Terre 68(Supplement):3–70.
Goloboff, P., J. C. Farris, and K. C. Nixon. 2008. TNT, a free program for
phylogenetic analysis. Cladistics 24:774–786.
Head, J. J. 1998. A new species of basal hadrosaurid (Dinosauria, Ornithis-
chia) from the Cenomanian of Texas. Journal of Vertebrate Paleon-
tology 18:718–738.
Head, J. J. 2001. A reanalysis of the phylogenetic position of Eolambia
caroljonesa (Dinosauria, Iguanodontia). Journal of Vertebrate Pale-
ontology 21:392–396.
Horner, J. R., D. B. Weishampel, and C. A. Forster. 2004. Hadrosauri-
dae; pp. 438–463 in D. B. Weishampel, P. Dodson, and H. Osm ´
olska
(eds.), The Dinosauria, second edition. University of California Press,
Berkeley, California.
Lambe, L. M. 1914. On Gryposaurus notabilis, a new genus and species
of trachodont dinosaur from the Belly River Formation of Alberta,
with a description of the skull of Chasmosaurus belli. The Ottawa
Naturalist 27:145–155.
Laurent, Y., L. Cavin, and M. Bilotte. 1999. D´
ecouverte d’un gisement
`
a vert´
ebr´
es dans le Maastrichtien sup´
erieur des Petites-Pyr´
en´
ees.
Comptes rendus de l’Acad´
emie des Sciences. S´
erie IIa, Sciences de
la Terre et des plan`
etes 328:781–787.
Leidy, J. 1858. Hadrosaurus foulkii, a new saurian from the Cretaceous
of New Jersey. Proceedings of the Academy of Natural Sciences of
Philadelphia 10:215–218.
L´
opez-Mart´
ınez, N., J. I. Canudo, L. Ard´
evol, X. Pereda-Suberbiola, X.
Orue-Etxebarria, G. Cuenca-Besc ´
os,J.I.Ruiz-Ome
˜
naca,X.Mure-
laga, and M. Feist. 2001. New dinosaur sites correlated with Upper
Maastrichtian pelagic deposits in the Spanish Pyrenees: implications
for the dinosaur extinction pattern in Europe. Cretaceous Research
22:41–61.
Lund, E. K., and T. A. Gates. 2006. A historical and biogeographical ex-
amination of hadrosaurian dinosaurs; pp. 263–276 in S. G. Lucas and
R. M. Sullivan (eds.), Late Cretaceous Vertebrates from the West-
ern Interior. New Mexico Museum of Natural History and Science
Bulletin 35.
Maddison, W. P., and D. R. Maddison. 2010. Mesquite: A Modular System
for Evolutionary Analysis, version 2.73. Available at mesquitepro-
ject.org. Accessed May 30, 2012.
Marsh, O. C. 1881. Principal characters of American Jurassic dinosaurs.
Part V. American Journal of Science, Series 3 21:417–423.
Maryanska, T., and H. Osm ´
olska. 1981. Postcranial anatomy of Saurolo-
phus angustirostris with comments on other hadrosaurs. Palaeontolo-
gia Polonica 46:119–141.
Nagao, T. 1936. Nipponosaurus sachalinensis, a new genus and species of
trachodont dinosaur from Japanese Saghalien. Journal of the Fac-
ulty of Science, Hokkaido Imperial University. Series 4, Geology and
Mineralogy 3:185–220.
Nopcsa, F. 1903. Telmatosaurus, new name for the dinosaur Limnosaurus.
Geological Magazine, Series 4 10:94–95.
Ogg, J. G., G. Ogg, and F. M. Gradstein. 2008. The Concise Geologic Time
Scale. Cambridge University Press, New York, vi +177 pp.
Oms, O., and J. I. Canudo. 2004. Dataci ´
on magnetoestratigr´
afica de los di-
nosaurios del Cret´
acico terminal (Maastrichtiense superior) de Ar´
en
(Huesca, Unidad Surpirenaica Central). Geo-Temas 6 5:51–54.
Ostrom, J. H. 1961. Cranial morphology of the hadrosaurian dinosaurs of
North America. Bulletin of the American Museum of Natural History
122:33–186.
Owen, R. 1842. Report on British fossil reptiles. Part II. Reports of the
British Association for the Advancement of Science 11:60–204.
Parks, W. A. 1922. Parasaurophus walkeri, a new genus and species of
crested trachodont dinosaur. University of Toronto Studies, Geolog-
ical Series 13:1–32.
Parks, W. A. 1923. Corythosaurus intermedius, a new species of trachodont
dinosaur. University of Toronto Studies, Geological Series 15:1–57.
Pereda-Suberbiola, X. 2009. Biogeographical affinities of Late Cre-
taceous continental tetrapods of Europe: a review, affinit´
es
pal´
eobiogeographiques des t´
etrapodes continentaux du Cr´
etac´
e
sup´
erieur d’Europe: une mise au point. Bulletin de la Soci´
et´
e
G´
eologique de France 180:57–71.
Pereda-Suberbiola, X., J. I. Canudo, P. Cruzado-Caballero, J. L. Barco,
N. L ´
opez-Mart´
ınez, O. Oms, and J. I. Ruiz-Ome ˜
naca. 2009. The last
hadrosaurid dinosaurs of Europe: a new lambeosaurine from the
uppermost Cretaceous of Ar´
en (Huesca, Spain). Comptes Rendus
Palevol 8:559–572.
Prieto-Marquez, A. 2007. Postcranial osteology of the hadrosaurid di-
nosaur Brachylophosaurus canadensis from the Late Cretaceous
Downloaded by [University of Zaragoza], [Miguel Moreno-Azanza] at 08:09 12 November 2013

1384 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 33, NO. 6, 2013
of Montana; pp. 91–116 in K. Carpenter (ed.), Horns and Beaks:
Ceratopsian and Ornithopod Dinosaurs. Indiana University Press,
Bloomington and Indianapolis, Indiana.
Prieto-Marquez, A. 2010. Global phylogeny of Hadrosauridae (Di-
nosauria: Ornithopoda) using parsimony and Bayesian methods. Zo-
ological Journal of the Linnean Society 159:435–502.
Prieto-Marquez, A., and G. C. Salinas. 2010. A re-evaluation of Secer-
nosaurus koerneri and Kritosaurus australis (Dinosauria, Hadrosauri-
dae) from the Late Cretaceous of Argentina. Journal of Vertebrate
Paleontology 30:813–837.
Prieto-Marquez, A., and J. R. Wagner. 2009. Pararhabododon isonensis
and Tsintaosaurus spinorhinus: a new clade of lambeosaurine
hadrosaurids from Eurasia. Cretaceous Research 30:1238–
1246.
Prieto-M´
arquez, A., R. Gaete, G. Rivas, A. Galobart, and M. Boada.
2006. Hadrosauroid dinosaurs from the Late Cretaceous of Spain:
Pararhabdodon isonensis revisited and Koutalisaurus kohlerorum,
gen. et sp. nov. Journal of Vertebrate Paleontology 26:929–
943.
Pu´
ertolas, E., J. I. Canudo, and P. Cruzado-Caballero, 2011. A new
crocodylian from the Late Maastrichtian of Spain: implications for
the initial radiation of crocodyloids. PLoS ONE 6:e20011.
Pujalte, V., and B. Schmitz. 2005. Revisi ´
on de la estratigraf´
ıa del Grupo
Tremp (“Garumniense”, Cuenca de Tremp-Graus, Pirineos merid-
ionales). Geogaceta 38:79–82.
Riera, V., O. Oms, R. Gaete, and A. Galobart. 2009. The end-Cretaceous
dinosaur succession in Europe: the Tremp Basin record (Spain).
Palaeogeography, Palaeoclimatology, Palaeoecology 283:160–
171.
Rozhdestvensky, A. K. 1966. [New iguanodonts from Central Asia.
Phylogenetic and taxonomic relationships between late Iguan-
odontidae and early Hadrosauridae]. Palaeontologicheskii Zhurnal
1966:103–116. [Russian; English translation: Rozhdestvensky, A. K.
1967. New iguanodonts from Central Asia. International Geology
Review 9:556–566].
Seeley, H. G. 1887. On the classification of the fossil animals commonly
called Dinosauria. Proceedings of the Royal Society of London
43:165–171.
Sereno, P. C. 1998. A rationale for phylogenetic definitions, with applica-
tion to the higher-level taxonomy of Dinosauria. Neues Jahrbuch f ¨
ur
Geologie und Pal¨
aontologie, Abhandlungen 210:41–83.
Sternberg, C. M. 1953. A new hadrosaur from the Oldman Formation of
Alberta: discussion of nomenclature. National Museum of Canada
Natural History Papers 128:1–12.
Sues, H.-D., and A. Averianov. 2009. A new basal hadrosauroid di-
nosaur from the Late Cretaceous of Uzbekistan and the early radi-
ation of duck-billed dinosaurs. Proceedings of the Royal Society B
267:2549–2555.
Suzuki, D., D. B. Weishampel, and N. Minoura. 2004. Nipponosaurus
sachalienensis (Dinosauria: Ornithopoda): anatomy and systematic
position within Hadrosauridae. Journal of Vertebrate Paleontology
24:145–164.
Walker, J. D., and J. W. Geissman (eds.). 2009. 2009 Geologic Time Scale.
Geological Society of America. doi: 10.1130/2009.CTS004R2C.
Weishampel, D. B., D. B. Norman, and D. Grigorescu. 1993. Tel-
matosaurus transsylvanicus from the Late Cretaceous of Romania:
the most basal hadrosaurid dinosaur. Palaeontology 36:361–385.
Williamson, T. E. 2000. Review of Hadrosauridae (Dinosauria, Ornithis-
chia) from the San Juan Basin, New Mexico; pp. 191–213 in S. G. Lu-
cas and A. B. Heckert (eds.), Dinosaurs of New Mexico. New Mexico
Museum of Natural History and Science Bulletin 17.
Submitted September 18, 2012; revisions received January 23, 2013;
accepted January 28, 2013.
Handling editor: You Hailu.
APPENDIX 1. Character coding scores for the 119 characters of
Godefroit et al. (2012), to which we have added one more charac-
ter from Prieto-M´
arquez (2010, character 82) in Arenysaurus and
Blasisaurus.
Arenysaurus
????? ????? ????? ???1? ?1?1? ?1??? ??111 10111 01?01 1100?
110?0 110?1 1111? ??1?1 11111 ??1?? ????? 101?? 1???? ??01?
????0 0111? ????? ???0
Blasisaurus
????? ????? ????? ????? ????? ?1211 0???? ????? ????? ?????
????0 11011 1111? ????? ????? ????? ????? ????? ????? ????1
????0 01??? ????? ???1
Downloaded by [University of Zaragoza], [Miguel Moreno-Azanza] at 08:09 12 November 2013