A New Species of Gryposaurus (Dinosauria - Hadrosauridae) From The Late Campanian Kaiparowits Formation, Southern Utah, USA

Zoological Journal of the Linnean Society, 2007, 151, 351–376.
With 16 figures
A new species of Gryposaurus (Dinosauria:

Hadrosauridae) from the late Campanian Kaiparowits
Formation, southern Utah, USA
TERRY A. GATES* and SCOTT D. SAMPSON
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Department of Geology and Geophysics and Utah Museum of Natural History, University of Utah,
1390 E. Presidents Circle, Salt Lake City, UT 84112, USA
Received March 2007; accepted for publication July 2007
A new species of the hadrosaurine hadrosaurid Gryposaurus was discovered in the late Campanian Kaiparowits
Formation of southern Utah. Gryposaurus monumentensis, sp. nov. is distinguished from other Gryposaurus
species by possessing a more robust skull, enlarged clover-shaped prongs on the predentary oral margin, an
anteroposteriorly narrow infratemporal fenestra, and other autapomorphies plausibly associated with feeding
adaptations. The derived morphology revealed in G. monumentensis necessitates revision of the generic diagnosis
of Gryposaurus, including the addition of synapomorphies that further aid in distinguishing this taxon from
Kritosaurus. A revised phylogenetic analysis places Gryposaurus within a monophyletic clade that includes
Brachylophosaurus and Maiasaura. Gryposaurus monumentensis represents the most southern example of
Gryposaurus, and underlines the remarkable diversification and long duration of this genus. Based on the
phylogenetic, geographical, and stratigraphic evidence at hand, Gryposaurus was the most diverse genus within
Hadrosaurinae; it also possessed one of the largest geographical and stratigraphic distributions, spanning more
than five million years of the Campanian, and ranging from Alberta in the north to Utah (and possibly Texas) in
the south. © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 151, 351–376.
ADDITIONAL KEYWORDS: biogeography – Cretaceous – dinosaur – hadrosaur – hadrosaurinae –

ornithopod.
INTRODUCTION Kritosaurus, and described the species K. incurvi-

manus, also from the Dinosaur Park Formation. Sub-
Gryposaurus is a hadrosaurine (noncrested or solid-
sequently, Lull & Wright (1942) assigned G. notabilis
crested) hadrosaurid distinguished by its deep skull
to the genus Kritosaurus, designated Gryposaurus as
and prominent nasal arch. The type species, Grypo-
a nomen dubium, and claimed that all hadrosaurids
saurus notabilis, was described by Lambe (1914) from
in which the infratemporal fenestra exceeds the orbit
the Dinosaur Park Formation of Alberta, Canada.
in area belong to the former genus. However, more
Four years earlier, Barnum Brown described a partial
recently, Horner (1992) outlined multiple distinctions
skull – missing the premaxillae and nasals – from the
between the two genera, including the shape of the
Kirtland Formation of New Mexico, which he named
frontal–nasal contact and morphology of the nasal
Kritosaurus navajovius (Brown, 1910). Both taxa
ornamentation. In addition, Horner resurrected
possess a deep skull with large dentaries and preden-
Gryposaurus, created the new combination of G. in-
taries and infratemporal fenestrae larger than the
curvimanus, and described yet another species,
orbit. On the basis of these similarities, Parks (1919)
G. latidens, characterized by relatively wider dentary
considered Gryposaurus to be a junior synonym of
teeth than in other species.
The Kaiparowits Formation is an 800-m-thick
package of rocks deposited between 76.1 and
*Corresponding author. E-mail: tgates@umnh.utah.edu 74.0 Mya on a wet, humid alluvial plain setting with
© 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 151, 351–376 351
352 T. A. GATES and S. D. SAMPSON
paludal and riparian environments (Roberts, Deino &

Chan, 2005; Roberts, 2007). Further, Roberts et al.
(2005) demonstrated that the Kaiparowits is tempo-
rally equivalent with many other fossiliferous forma-
tions within the Western Interior Basin – most
significantly for this discussion, the Dinosaur Park
Formation, where G. notabilis and G. incurvimanus
co-occur. The Kaiparowits Basin Project (KBP),
launched in 2000 by the University of Utah, is a
multi-institution endeavour that has focused on the
unravelling of the macrovertebrate fossil record pre-

served within the Kaiparowits Formation. To date,
KBP has met with considerable success (Sampson
et al., 2004), resulting in the discovery of several new
species, including a new oviraptorosaur (Zanno &
Sampson, 2005), a chasmosaurine ceratopsid (Smith
et al., 2004), and a hadrosaurid (this paper).
Hadrosaurid fossils are extremely abundant in the
Kaiparowits Formation. Yet, the only taxon described
to date from this unit is the lambeosaurine (crested
hadrosaurid) Parasaurolophus cf. cyrtocristatus
(Weishampel & Jenson, 1979; Sullivan & Williamson,
1999; Gates, 2004), known from multiple partial
skulls and a partial postcranium (T. A. Gates; E. K.
Lund; M. A. Getty; J. I. Kirkland; A. L. Titus; D.
Deblieux; C. A. Boyd & S. D. Sampson, unpubl. data).
Previous hadrosaurine materials collected from the
Kaiparowits Formation have generally been placed
into Kritosaurus, which has become a waste-basket
taxon for most nondiagnostic hadrosaurine materials Figure 1. Lateral view of type specimen of Gryposaurus
collected in the southern region of the Western Inte- monumentensis gen. et sp. nov. skull, RAM 6797: A,
rior Basin. However, in August 2004, personnel from photograph of left side of skull; B, line drawing of skull
based on photograph in (A) and showing slight reconstruc-
the Raymond M. Alf Museum of Palaeontology exca-
tion. Ar, articular; D, dentary; F, frontal; J, jugal; La,
vated a mostly complete hadrosaurine skull from the
lacrimal; Mx, maxilla; Na, nasal; Pd, predentary; Pf, pre-
Kaiparowits Formation. This skull and additional
frontal; Pm, premaxilla; Po, postorbital; Qj, quadratojugal;
specimens collected by field crews from the Utah
Qu, quadrate; Sq, squamosal; Su, surangular. Scale bar,
Museum of Natural History are clearly referable to 10 cm.
Gryposaurus, and represent yet another new species
(Fig. 1). This paper describes the new taxon, revises
the generic diagnosis of Gryposaurus, and discusses Ontario Museum, Toronto, ON, Canada; TMP, Royal
the phylogenetic, palaeobiological, and biogeographi- Tyrrell Museum of Palaeontology, Drumheller, AB,
cal significance of this dinosaur. In addition, biostrati- Canada; UMNH, Utah Museum of Natural History,
graphic patterns are revealed in the Kaiparowits University of Utah, Salt Lake City, UT, USA.
Formation through comparison of the temporal span
of an unidentified, undescribed specimen of Gryposau-
SYSTEMATIC PALAEONTOLOGY
rus in the base of the formation with the temporal
occurrences of the new Gryposaurus taxon described DINOSAURIA OWEN, 1842
herein. ORNITHISCHIA SEELEY, 1887
HADROSAURIDAE COPE, 1869
INSTITUTIONAL ABBREVIATIONS HADROSAURINAE COPE, 1869
AMNH, American Museum of Natural History, New GRYPOSAURUS LAMBE, 1914
York, NY, USA; CMN, Canadian Museum of Nature, Etymology: Derived from the Latin for ‘hooked beak’,
Ottawa, ON, Canada; OTM, Old Trail Museum, intended by Lambe to reference the arch on the dorsal
Choteau, MT, USA; RAM, Raymond M. Alf Museum surface of the nasal, resembling a gryphin, and the
of Palaeontology, Claremont, CA, USA; ROM, Royal Greek ‘saurus’ for lizard.
© 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 151, 351–376
NEW SPECIES OF GRYPOSAURUS 353
Revised diagnosis: Hadrosaurine hadrosaurids pos- other species. Gryposaurus incurvimanus also pos-
sessing the following autapomorphies: dorsolateral sesses an anteroposteriorly shortened skull relative to
flare on the medial margin of the premaxillary the maximum skull depth.
lateral process; unexcavated nasal arch located ante-
rior to the orbits; and sigmoidal nasofrontal suture Holotype: ROM 764, mostly complete skull and com-
characterized by a small median process of the nasal plete postcranium.
inserting between the midline of the frontals.
Members of the genus also possess the following
unique combination of characters: dorsoventrally
deep skull; anterior process of nasal terminates along GRYPOSAURUS LATIDENS HORNER, 1992
the dorsal margin of the external naris; ventral nasal Diagnosis (after Horner, 1992): Dentary teeth shorter

process comprising approximately 25% of the ventral and wider than in other species of the genus. In
margin of the external nares; abbreviated circum- addition, G. latidens possesses the following unique
narial depression, extending posteriorly only as far combination of characters: arched lateroventral
as the lacrimal; jugal with large offset posteroventral margin on the maxilla; nasal arch positioned near the
flange, postorbital process at or near 90°, and narrow orbits and rising above the level of the frontals; and
posterior process; and frontal contribution to the broad ‘U’-shaped posterior margin of the external
orbital rim. naris. The humerus of G. latidens is larger and more
robust than that of G. incurvimanus.
GRYPOSAURUS NOTABILIS LAMBE, 1914 Holotype: AMNH 5465, partial skull and nearly com-
plete skeleton.
Revised diagnosis: Hadrosaurids of the genus Grypo-
saurus with the following unique combination of char-
acters: sigmoidal lateroventral margin of the maxilla;
large nasal arch located near the orbits and rising GRYPOSAURUS MONUMENTENSIS SP. NOV.
above the level of the frontals; narrow ‘U’-shaped
Etymology: In reference to the Grand Staircase-
posterior margin of the external naris; squamosal
Escalante National Monument in southern Utah,
posterodorsal margin well above the level of the skull
where all referred materials of this species have been
roof; infratemporal fenestra larger than the orbit; and
recovered.
rounded mandibular foramen. The skull of G. notabi-
lis is generally more robust than that of G. latidens
Diagnosis: Hadrosaurids of the genus Gryposaurus
and G. incurvimanus, but less so than that of
possessing: anterodorsal process of maxilla seen
G. monumentensis.
through the external nares; subparallel anterior and
posterior margins of the infratemporal fenestra;
Holotype: CMN 2278, complete articulated skull.
anteroposterior width of the infratemporal fenestra
approximately half of the width of the orbit; preden-
tary with large, pronged denticles; anterior portion of
GRYPOSAURUS INCURVIMANUS PARKS, 1919 the dentary sharply downturned; surangular with
Revised diagnosis: Hadrosaurids of the genus Grypo- distinct dorsal process on the median ridge of suran-
saurus possessing: dorsal premaxillary process that is gular; and ovoid mandibular foramen. The following
more concave posteriorly than in other species of the suite of characters is also present: sigmoidal lat-
genus; nasal arch smaller than in other species and eroventral margin of the maxilla; large nasal arch
positioned well anterior to the orbit and rising to a located near the orbits rising above the level of the
level below the frontals; jugal with a small spur on frontals; narrow ‘U’-shaped posterior margin of the
the posterior margin of the posteroventral flange; and external naris; and squamosal raised well above
slightly excavated ventral surface of the nasal hump. the level of the skull roof. Gryposaurus monumenten-
In addition, G. incurvimanus possesses the following sis is significantly more robust than any other species
unique combination of characters: nearly straight of Gryposaurus. It is most similar to G. notabilis, but
lateroventral margin of the maxilla; infratemporal differs in having more steeply angled premaxillae,
fenestra larger than the orbit; and broad ‘U’-shaped mediolaterally wider dentary, and generally more
posterior margin of the external naris. Gryposaurus prominent jugal tubercle.
incurvimanus is significantly more gracile than either
G. notabilis or G. monumentensis. It has a smaller Holotype: RAM 6797, largely articulated skull
premaxillary lip than the other species, as well as lacking most of the right side of the skull (Figs 1, 3,
premaxillae that are angled more steeply than in 5A, 7A, B, 9, 11, 12, 13A, 14, 15).

Figure 2. Map showing location of Grand Staircase-Escalante National Monument (GSENM) within Utah.
Referred materials: UMNH VP 13970 (Figs 4, 7C, D, G. monumentensis (UMNH VP 13970) preserves
8, 13B), partial, disarticulated subadult skull that many delicate elements, such as the vomers and
includes both maxillae and dentaries, right nasal, complete nasals. All elements of the craniofacial skel-
partial quadrate, right jugal, right vomer, right pre- eton, except the parietals and frontals, are preserved
frontal; UMNH VP 12265, partial skull and skeleton in detail on the type specimen and/or UMNH VP
preserving both maxillae, left dentary, left quadrate, 13970.
and partial jugal, left humerus, right and left scapu-
lae, right coracoid, multiple ribs, both ilia, pubes, and
ischia, most of the dorsal, sacral, and caudal series, CRANIUM
including skin impressions on the right side of the General: Overall, the skull of G. monumentensis is
vertebral series. extremely robust, much more so than that of any
other species of the genus. It is also relatively large
Horizon: All known specimens are from the middle and compact – that is, relatively deep dorsoventrally
unit of the Kaiparowits Formation (Fig. 2; Upper Cre- and abbreviated anteroposteriorly. This combination
taceous: late Campanian). of increased robustness and anteroposterior shorten-
ing results in subtle, but nonetheless significant,
modifications to much of the craniofacial skeleton
DESCRIPTION when compared with other species of Gryposaurus.
The G. monumentensis holotype specimen, RAM
6797, was found in an ancient point bar deposit in the Premaxilla: The premaxilla of G. monumentensis is
middle unit of the Kaiparowits Formation. The left robust and steeply angled, giving the anterodorsal
half of the skull sank into soft sediment, remaining region of the skull its characteristic shape (Fig. 1). It
completely articulated, and is preserved in relatively contacts the maxilla and lacrimal posterolaterally
pristine condition. The right side of the skull suffered along its lateral process and the nasal through both
from disarticulation and loss of elements, apparently the dorsal and lateral processes (Fig. 1). For the pur-
the result of decay and transport in the river current; poses of description, this element is divided into three
evidence of fluvial influence was seen in the right regions: (1) an anterior region; (2) a dorsal process;
quadrate, which was recovered on top of the right and (3) a lateral process.
maxilla. The frontals, parietals, right side of the The oral margin is highly rugose (Fig. 3). In addi-
braincase, and posterior nasals were partially eroded tion, a large patch of rugose bone is present on the
prior to excavation, preventing detailed examination midline at the anterior limit of the interpremaxillary
of these elements. The nasals, however, do preserve contact, a feature not observed in any other Grypo-
the nasal ornamentation, although not in articula- saurus species. It is likely that the highly rugose oral
tion. A disarticulated subadult specimen referred to margin and large midline rugosity participated in

Figure 4. Lateral view of Gryposaurus monumenten-
sis gen. et sp. nov. left nasal, UMNH VP 13970. en,
external naris; na, nasal arch; nap, nasal anterior process;
nfs, nasofrontal suture. Scale bar, 5 cm.
Figure 3. Lateral view of Gryposaurus monumenten-

sis gen. et sp. nov. left premaxilla, RAM 6797. en, exter- bilis (ROM 873), and significantly more so than that
nal nares; MX, maxilla; or, oral margin rugosities; pmd, of G. incurvimanus (TMP 80.22.1) and G. latidens
dorsal process; pmf, premaxillary foramen; pml, lateral (MOR 553s-7-18-91-107). Despite differences in
process; pmlf, premaxilla lateral process flare; pmlp, pre- robustness and width, all Gryposaurus species share
maxillary lip; pms, premaxillary shelf. Scale bar, 5 cm. an upturned premaxillary lip and dorsal flaring of the
lateral process.
anchoring a similarly robust keratinous rhamphoth- Nasal: The nasal is an axe-shaped element in lateral
eca (Morris, 1970). In dorsal view, the oral margin view that contacts anteriorly with the premaxilla,
expands laterally to a maximum width at least equal posteroventrally with the premaxilla, lacrimal, and
to the width of the skull at the postorbital. Unlike prefrontal, and posteriorly with the frontals (Fig. 1).
Brachylophosaurus (Prieto-Marquez, 2005), the pre- The anterior process of the nasal flanks the premax-
maxillary lip is upturned and slightly folded posteri- illary dorsal process laterally, terminating prior to the
orly (Fig. 3), a condition that is less pronounced in anterior margin of the external nares, a condition
Gryposaurus than in either Prosaurolophus (Horner, shared with the other species of Gryposaurus, Maia-
1992) or Edmontosaurus (Lambe, 1920). Posterior to saura (Horner, 1983), Bactrosaurus (Godefroit et al.,
the premaxillary lip, the premaxillary shelf is broad 1998), Telmatosaurus (Weishampel, Norman & Grig-
and bulges dorsally as a result of a concave depres- orescu, 1993), and most other basal iguanodontians
sion on the ventral side of the element. The large (Norman, 2004).
premaxillary foramen resides posteroventral to the The transversely compressed nasal arch is the most
dorsal process (Fig. 3). distinctive characteristic distinguishing Gryposaurus
In lateral view, the dorsal process of the premaxilla from all other hadrosaurids (Figs 1, 4). Gryposaurus
arches posterodorsally more than in any other incurvimanus (TMP 80.22.1) exhibits a smaller arch
hadrosaurine taxon except G. incurvimanus (TMP positioned more anteriorly than that in G. notabilis
80.22.1; the premaxillae are unknown for Kritosau- (CMN 2278, ROM 873), G. latidens (AMNH 5465,
rus). The dorsal process is approximately 75% of the MOR 478), and G. monumentensis (RAM 6797,
length of the lateral process, continuing almost to the UMNH VP 13970; Fig. 5).
posterior extent of the external nares. It bears a Posteriorly, the nasals contact the frontals in a
shallow lateral groove to secure the nasal anterior sigmoidal suture resulting from small, paired pro-
process. cesses of the nasals that insert along the midline of
The lateral process of the premaxilla extends pos- slightly divergent frontals (Horner, 1992; Fig. 6). The
terodorsally to rest on the lacrimal behind the exter- frontal–nasal contact is not preserved on the type
nal nares. Unique amongst other hadrosaurid genera, specimen of G. monumentensis. However, the poste-
the medial margin of the premaxilla lateral process rior portion of the right nasal in UMNH VP 13970
flares sharply dorsally approximately midway along indicates that this sigmoidal suture was probably
the ventral margin of the external naris (Fig. 3). In present in the Utah taxon (Fig. 4). The characteristic
comparison with other Gryposaurus species, the pre- sigmoidal nasofrontal suture differs from the posteri-
maxilla of G. monumentensis (RAM 6797) is slightly orly concave suture of Brachylophosaurus (Prieto-
more robust than that of some specimens of G. nota- Marquez, 2005), and the posteriorly lobate suture of
Figure 5. Line drawings of the four Gryposaurus species

in lateral view, showing synapomorphies of the genus and
other comparative features: A, G. monumentensis gen.
et sp. nov. (RAM 6797); B, G. notabilis (ROM 873); C,
G. incurvimanus (TMP 80.22.1); D, G. latidens (after
Horner, 1992; no scale included). itf, infratemporal fenes-
tra; mf, mandibular foramen; na, nasal arch; pmlf, lateral
process flare. Scale bar, 10 cm.
䉳

Figure 6. Dorsal view of frontal–nasal suture in Grypo-
saurus incurvimanus (ROM 764). F, frontal; Na, nasal; nfs,
nasofrontal suture; Pa, parietal; Pf, prefrontal; Po, postor-
bital. Scale bar, 5 cm.
Prosaurolophus and Naashoibitosaurus (NMMNH

P-16106) that inserts a small frontal process between
splayed nasals (Horner, 1992).
In lateral view, the posteroventral region of the
nasal forms the axe head of the element (Figs 1, 4).
The circumnarial depression (Hopson, 1975) is
expressed as a shallow halo around the external
nares. In contrast, the circumnarial depression
extends much further posteriorly in several other
hadrosaurines, terminating just anterior to the orbits
[Anasazisaurus (BYU 12950), Naashoibitosaurus
(NMMNH P-16106); Horner, 1992] or even on the
skull roof (Prosaurolophus blackfeetensis and Sau-
rolophus angustirostris; Maryanska & Osmólska,
1981; Horner, 1992).
Not surprisingly, the G. monumentensis nasal
underwent a shape change during ontogeny, as evi-
denced by comparison of nasals from the type speci-
men and the subadult UMNH VP 13970. In general,

Figure 7. Maxillae from Gryposaurus monumentensis gen. et sp. nov., specimens RAM 6797 and UMNH VP 13970:
A, left maxilla in lateral view with surrounding elements from RAM 6797; B, posterior region of right maxilla from UMNH
VP 13970; C, right maxilla from RAM 6797 in lateral view; D, anterior region of left maxilla from UMNH VP 13970. D,
dentary; ecr, ectopterygoid ridge; ecs, ectopterygoid shelf; J, jugal; jp, jugal process; La, lacrimal; madp, anterodorsal
process; mavp, anteroventral process; mdp, maxilla dorsal process; mf, maxillary foramen; Mx, maxilla; nf, nutrient
foramen; palp, palatine process; Pm, premaxilla; ptp, pterygoid process. Scale bar, 5 cm.
it appears that this species increased the size and Brachylophosaurus (Prieto-Marquez, 2005) and Maia-
prominence of the nasal arch and decreased the angle saura (Horner, 1983).
of the anterior process. A rounded dorsal process (Fig. 7A) rises steeply
from the central region of the maxilla. The structure
Maxilla: The robust maxilla of G. monumentensis can is broader anteroposteriorly than in other hadrosau-
be subdivided into three regions: an anterior region rine taxa, being most similar to G. notabilis (CMN
that includes a pair of premaxillary processes; a 2278). The largest maxillary foramen opens at the
central region contacting the jugal and lacrimal; and base of the dorsal process (Fig. 7A, D). The dorsal
a posterior region that unites the palate with the process contacts the jugal along a broad, sigmoidal
facial skeleton (Fig. 7). Two large, anteriorly directed suture (Fig. 7A, C). The large overhanging jugal
processes extend from the anterior-most edge of the process is larger than that in Brachylophosaurus
maxilla (Fig. 7C, D). The anteroventral process is (MOR 1071 8-13-98-559) and in other species of
short and broad in order to support the lateral process Gryposaurus, but is more similar in size to that in
of the premaxilla dorsally. The longer anterodorsal P. blackfeetensis (Horner, 1992).
process displays a medial expansion comparable in Contact with the palatine, ectopterygoid, and ptery-
size with that of Naashoibitosaurus (NMMNH goid (Fig. 7B, C) occurs on the posterior region of the
P-16106), but larger than that of P. blackfeetensis maxilla. As exposed laterally on the holotype, the
(Horner, 1992). In G. monumentensis, the anterodor- posterior region is narrow dorsoventrally, because of
sal process is visible within the external naris when an expanded ventral tubercle of the jugal (see below).
the skull is viewed laterally (Fig. 1). This contrasts This dorsoventral compression is present, although
with all other species of Gryposaurus, but is similar to not as developed, in G. notabilis (ROM 873), whereas
contacts the posterior side of the maxilla dorsal

process (Fig. 8; Maiasaura has a straight ventral
margin; see Horner, 1990). Edmontosaurus regalis
(e.g. CMN 2288), P. maximus (e.g. TMP 84-1-1), and
Saurolophus osborni (e.g. AMNH 5220) all possess
relatively convex ventral margins of the jugal anterior
process. In G. monumentensis, the ventral tubercle of
the anterior process is more hypertrophied than in
any other species of Gryposaurus, and more so than in
hadrosaurids generally, extending ventrally almost to
the alveolar margin. However, the size and shape of

this feature vary significantly in individuals of G. no-
tabilis (e.g. ROM 873 and CMN 2278), and should be
used with caution in a phylogenetic context.
Figure 8. Lateral view of juvenile right jugal from
The postorbital process ascends at nearly a 90°
UMNH VP 13970. jap, jugal anterior process; jpp, jugal angle (Fig. 8). Maiasaura, Brachylophosaurus, Krito-
posterior process; pop, postorbital process; pvf, posteroven- saurus, Anasazisaurus, and Naashoibitosaurus share
tral flange. Scale bar, 5 cm. a nearly vertical postorbital process, whereas the
process of Prosaurolophus spp. and Saurolophus spp.
is slightly angled posteriorly, and steeply angled in
Edmontosaurus (Brown, 1910; Lambe, 1920; Horner,
the posterior maxilla has a significantly broader expo-
1983, 1992; Prieto-Marquez, 2005; Lucas et al., 2006).
sure in G. incurvimanus (TMP 80.22.1) and G. lati-
In G. monumentensis (RAM 6797 and UMNH VP
dens (Horner, 1992; Fig. 5). A long, broad palatine
13970), and Gryposaurus generally, the posteroven-
process and triangular pterygoid process emanate
tral flange is a moderately sized extension of the jugal
posterior to the dorsal process along the tall median
(Fig. 8). Gryposaurus incurvimanus (TMP 80.22.1)
ridge of the posterior region (Fig. 7B, C). The ectop-
bears a small spur on the posterior edge of the flange,
terygoid closely adheres to the broad ectopterygoid
which is probably an apomorphy of the species. The
shelf, draping over the rounded posterior end of the
prominent flange of Brachylophosaurus (Prieto-
maxilla (Fig. 7C). The ectopterygoid ridge (Fig. 7B)
Marquez, 2005) and Maiasaura (Horner, 1983) is only
is not as sharply developed as in P. blackfeetensis
slightly more exaggerated than that of Gryposaurus,
(Horner, 1992) or Naashoibitosaurus (NMMNH
but is quite distinct from the shallow flange of
P-16106). There are approximately 49 tooth positions
P. blackfeetensis (Horner, 1992).
along a sinusoidal tooth row. The lateroventral
The posterior process is long and slender, again
margin of the maxilla in G. monumentensis (RAM
differing from that of both P. blackfeetensis (Horner,
6797) is dramatically sigmoidal, as in G. notabilis
1992) and Brachylophosaurus (Prieto-Marquez,
(ROM 873, CMN 2278), yet differs from the nearly
2005), which possess long and broad or short and
straight condition in G. incurvimanus (TMP 80.22.1,
expanding processes, respectively.
and possibly ROM 764) and the slightly arching con-
In UMNH VP 13970, the posteroventral flange is
dition in G. latidens (Horner, 1992; Fig. 5) and most
smaller relative to the total size of the jugal than in
other hadrosaurine taxa. An arching line of the dental
RAM 6797 (Fig. 1), suggesting positive allometry
foramina is present on the medial side of the maxilla
within this structure. The postorbital process likewise
(Fig. 7C).
changes slightly through ontogeny in that the angle of
ascent morphs from about 60° in UMNH VP 13970 to
Jugal: The jugal (Fig. 8) contacts the maxilla, the nearly vertical in RAM 6797, ultimately decreasing
lacrimal, the postorbital, the quadrate, and quadra- the size of the orbit and increasing the size of the
tojugal (Fig. 1). Overall, this element is most similar infratemporal fenestra with increasing skull length.
to that of G. notabilis (ROM 873, CMN 2278; Fig. 5),
although it shows distinct similarities with the jugal Lacrimal: As in other hadrosaurids, the lacrimal is a
of Kritosaurus (AMNH 5799, BYU 12950, NMMNH triangular bone that forms the anterior margin of the
P-16106; Brown, 1910; Lucas et al., 2006). orbit (Fig. 1). The dorsal surface of the lacrimal is
Unlike the anterior process of lambeosaurines, this sandwiched between the lateral process of the pre-
feature tends to be notably asymmetrical in hadro- maxilla laterally and the nasal medially. The prefron-
saurines. Specifically, in Gryposaurus, Brachylopho- tal contributes to the posterodorsal corner of the
saurus, and Kritosaurus, the ventral half of the jugal lacrimal, covering a small section of the lateral
head forms a large sweeping sigmoidal curve that surface. Contact between these two elements is
rosaurine taxa in that it is subtriangular in general

shape. Generally, the quadratojugal of the Utah taxon
is most comparable with that of Prosaurolophus
(Horner, 1992) and least similar to that of Edmonto-
saurus (Lambe, 1920). Posteriorly, a cupped region
accepts the lower portion of the quadrate lateral wing
and overlaps the quadratojugal notch on the quad-
rate. The posteroventral flange of the quadratojugal
appears more pronounced in G. monumentensis than
in other taxa.

Prefrontal: The prefrontal of G. monumentensis
UMNH VP 13970 is virtually identical to that of
G. latidens (MOR 553M 6-27-9-55), contacting with
the lacrimal anteroventrally (Fig. 1), the nasal medi-
ally, and the frontal medioposteriorly. It forms the
anterodorsal corner of the orbital rim. Two foramina
are situated near the junction of the smooth antero-
lateral margin and the heavily rugose posterolateral
margin. A large triangular groove on the anteroven-
tral surface of the prefrontal receives the lacrimal.
The medial region bows slightly dorsally and overlaps
the nasal. Posteriorly, the prefrontal has a large
process that inserts into an anterolateral depression
in the frontal.
Frontal: The morphology of the frontal is conserved

across known species of Gryposaurus, being broad,
depressed medially, and sculpted anteriorly with a
sigmoidal nasofrontal suture (Fig. 6). However, the
frontal is not well preserved on any specimens
Figure 9. Posterior view of left lacrimal–jugal contact in assigned to G. monumentensis, including the holo-
Gryposaurus monumentensis gen. et sp. nov. (RAM type. Only the lateral-most portion of the left frontal
6797). The left side of the figure is lateral and the right is preserved in RAM 6797, where it participates in
side is medial. Note the interlocking relationship between the formation of the orbital rim between the prefron-
these elements. J, jugal; La, lacrimal; lf, lacrimal foramen. tal and the postorbital (Fig. 1). This trait is present
in Gryposaurus, Brachylophosaurus, Telmatosaurus,
Bactrosaurus, and, indeed, most iguanodontians
accomplished through a triangular process on the (Norman, 1986; Weishampel et al., 1993; Godefroit
dorsal-most corner of the lacrimal that inserts into a et al., 1998; Prieto-Marquez, 2005).
receptive groove on the prefrontal. In posterior
aspect, the large, ovoid lacrimal foramen penetrates Postorbital: The postorbital possesses three distinct
anteriorly through the body of the element (Fig. 9). processes (anterior, squamosal, and jugal) contacting
Ventral to the lacrimal foramen, the jugal–lacrimal the frontal, squamosal, and jugal, respectively
articulation is an ‘S’-shaped facet that tightly bonds (Fig. 1). The squamosal process extends posteriorly,
the two elements together (Fig. 9). The ventral but is relatively shorter in G. monumentensis than in
margin of the lacrimal rests subhorizontally within G. notabilis (ROM 873 and CMN 2278) and G. incur-
the lacrimal groove on the maxilla dorsal process, vimanus (TMP 80.22.1 and ROM 764). The relative
closer in form to Edmontosaurus (Lambe, 1920) and shortening of the squamosal process contributes to
Naashoibitosaurus (NMMNH P-16106) than to the anteroposterior narrowing of the infratemporal
Brachylophosaurus (Prieto-Marquez, 2005) or Prosau- fenestra and supratemporal fenestra (Fig. 1; see
rolophus (Horner, 1992). below). Contact with the parietals occurs on a small
area of the postorbital medial to the base of the
Quadratojugal: Contacting with the jugal anteriorly squamosal process. A large pocket located on the
and the quadrate posteriorly (Fig. 1), the quadratoju- medial side of the postorbital, ventral to the contact
gal of G. monumentensis resembles that of other had- facet for the parietal, marks the insertion of the
feetensis (Horner, 1992). Posterior to the quadrate

cotylus, the postcotyloid process abuts the exoccipitals
along its posterior side and descends nearly the entire
length of the exoccipitals (Fig. 10). The medial region
of the squamosals is missing from RAM 6797,
although in the Gryposaurus sp. skull, UMNH VP
16669, as well as other species of Gryposaurus, the
medial rami are separated from one another along the
midline by a small ridge of parietal.
BRAINCASE

General: The left side of the braincase of RAM 6797
is only partially preserved, lacking the parietals, fron-
tals, supraoccipital, orbitosphenoids, and presphe-
noids. The following description is based on this
partial specimen.
Figure 10. Left squamosal from Gryposaurus sp. UMNH
VP 16669 in lateral view. poc, postcotyloid process; pog,
Fused exoccipital–opisthotic: The exoccipital makes
postorbital groove; prc, precotyloid process; prf, precotyloid
up the posterodorsal limit of the skull. Only a portion
fossa; qc, quadrate cotylus. Scale bar, 5 cm.
of the left exoccipital is preserved in the type speci-
men. It is shown in this specimen to contact the
squamosal, basioccipital, and pro-otic through two
laterosphenoid. The lateral margin (i.e. the postero- main regions: the paroccipital process and the exoc-
dorsal orbital rim) is highly rugose along its entire cipital base. The ventrolaterally hooking, pendant
length. paroccipital processes are relatively elongate, termi-
nating level with the base of the exoccipital, as in
Squamosal: The squamosal makes up the posterolat- Prosaurolophus (Horner, 1992), and shorter than in
eral section of the skull roof contacting the quadrate, Brachylophosaurus (MOR 1071). The paroccipital
postorbital, parietal, supraoccipital, and exoccipital process tightly abuts the postcotyloid process of the
(Fig. 1). Anteriorly, the postorbital process of the squamosal anteriorly. The entire median section of
squamosal terminates near the main body of the the left exoccipital is missing in RAM 6797, giving no
postorbital. A deep fossa occurs between the postor- indication of the morphology of this region. Unlike
bital process and the nearly vertical precotyloid Prosaurolophus (Horner, 1992), the exoccipitals do
process (Fig. 10). The precotyloid fossa is limited pos- not make up the ventral border of the foramen
terodorsally by a pronounced depression, but opens magnum, but are instead separated by a thin region
anteroventrally. This feature is present in Naashoi- of basioccipital. The basioccipital forms a long antero-
bitosaurus (NMMNH P-16106), Kritosaurus (AMNH posteriorly orientated articulation with the ventral
5799), Prosaurolophus (e.g. TMP 84.1.1 and Horner, processes of the exoccipital base. Although the
1992), and Brachylophosaurus (MOR 1071), but is sutures are obscured in RAM 6797, there seems to be
shallower and narrower in the last two taxa. The no evidence of the anterior projection of the exoccipi-
development of this fossa may be a result, in part, of tal that touches the basisphenoid in Prosaurolophus
the size of the precotyloid process, as Gryposaurus (Horner, 1992). Foramina for cranial nerves (CN) X,
has a larger process than Brachylophosaurus and XI, and XII are located on the lateral surface of the
Prosaurolophus – which possess smaller processes exoccipital base, with XII positioned posteriorly and X
and fossae – and Edmontosaurus (e.g. CMN 2288), in and XI sharing a more anterior opening. The poste-
which the precotyloid process is small, and does not rior border of CN VIII is composed of the anterior
possess a corresponding fossa. Within Gryposaurus, margin of the exoccipital, which contacts the pro-otic
the fossa appears deeper in G. monumentensis than in anteriorly.
G. notabilis (ROM 873) or G. incurvimanus (ROM
764, TMP 80.22.1), but is comparable with that of the Basioccipital: The basioccipital contacts the exoccipi-
G. notabilis holotype (CMN 2278). The quadrate tals posterodorsally, the pro-otic dorsally, and the
cotylus is squared off in a manner similar to that basisphenoid anteriorly. In posterior view, the basio-
in Brachylophosaurus (Prieto-Marquez, 2005). The ccipital is crescentic and dorsoventrally grooved. The
dorsal margin of the cotylus is nearly horizontal and dorsal margin of the crescent represents the base of
lacks the posterodorsal projection seen in P. black- the foramen magnum. In ventral aspect, the posterior
Pro-otic: The nearly pentagonal pro-otic contacts the

supraoccipital dorsally, the exoccipital posteriorly, the
basioccipital posteroventrally, and the basisphenoid
anteroventrally. The pro-otic contacts the laterosphe-
noid along its entire anterior border, forming the
posterior margin of the foramen for CN V (Fig. 11B).
A thin ridge separates CN V foramen from the more
ventral CN VII exit. In medial view, the CN VII
foramen shares an internal depression with that for
CN VIII, although the CN VIII opening occupies a
more posteroventral position within this depression

than is found in Prosaurolophus (Horner, 1992). The
anterior border for the external opening of CN VIII is
formed by the pro-otic.
Laterosphenoid: In anterior view, the laterosphenoid

is triradiate, possessing a ventral prong that articu-
lates with the basisphenoid and the opposite lat-
erosphenoid, a dorsomedial prong that contacts the
parietal, and a blunt-headed dorsolateral process that
inserts into a medial pocket on the postorbital. The
orbitosphenoid and frontal are sutured to the lat-
erosphenoid on its dorsal surface. The opening for CN
II is prominent on the ventral half of the element. The
remainder of the anterior region is not preserved.
Posterolaterally, the laterosphenoid creates the ante-
rior border of CN V (Fig. 11B), as in more derived
hadrosaurids, whereas the basisphenoid contributes
to the ventral border of CN V in at least Iguanodon
(Norman, 1980, 1986), Bactrosaurus (Godefroit et al.,
1998), and Telmatosaurus (Weishampel et al., 1993).
This conformation appears to represent a previously
unrecognized synapomorphy of Euhadrosauria (sensu
Weishampel et al., 1993) or Hadrosauridae (sensu
Forster, 1997). A large, anterolaterally orientated
depression traces the path of CN V to the lateral edge
of this element in G. monumentensis (RAM 6797). A
Figure 11. Gryposaurus monumentensis gen. et sp. long extension of the laterosphenoid encircles the
nov. (RAM 6797) braincase: A, ventral view; B, anterior dorsal margin of CN V, overlapping the pro-otic
region in lateral view. bpt, basipterygoid process; Bs, (Fig. 11B). The extension is long and tapered, termi-
basioccipital; bsmm, basioccipital median mound; Bsp, nating well past the apex of the opening for CN V,
basisphenoid; bst, basitubera; btr, basisphenoid transverse which differs substantially from the relatively short,
ridge; CN, cranial nerve; Exo, exoccipital; Ls, laterosphe- broad extension in P. blackfeetensis (Horner, 1992).
noid; mvp, median ventral process; Pr, pro-otic; psp, paras- The morphology of the posterior extension in Brachy-
phenoid process; Qu, quadrate. Scale bar, 5 cm.
lophosaurus (MOR 1071) is more similar to that of
Gryposaurus, except that it extends further around
region of the basioccipital is convex, whereas the the foramen for CN V in the former taxon. Ventral to
anterior region is dominated by the laterally posi- the CN V opening is a raised, slightly concave plateau
tioned paired basitubera (Fig. 11A). They are low, that resembles a structure seen in Prosaurolophus
bulbous, and distinctly separated from the posterior (Horner, 1992); however, in RAM 6797, it is slightly
half of the basioccipital by a large groove not seen in taller with sharper and more distinct edges.
Brachylophosaurus (MOR 1071). A much smaller
mound is present medial to the basitubera along the Basisphenoid: The basisphenoid contacts the basioc-
midline. Overall, this configuration closely resembles cipital posteriorly in a long broad facet (Fig. 11A). In
that of other hadrosaurines, such as Prosaurolophus addition, this element contacts the pro-otic postero-
(Horner, 1992). dorsally, laterosphenoid anterodorsally, and pterygoid
is composed of a large lateral condyle that contacts

the surangular, and a somewhat smaller, dorsomedi-
ally positioned condyle that contacts the articular.
In lateral view, the quadrate lateral wing expands
anteroventrally in a gentle arch, but is interrupted
two-thirds along the length of the shaft by a tall
quadratojugal notch (Fig. 12). The quadratojugal
notch is asymmetrical and subtriangular, with a
dorsal margin descending posterodorsally at approxi-
mately 60° and a short subhorizontal ventral margin.
Brachylophosaurus, on the other hand, possesses a

nearly symmetrical quadratojugal notch (Prieto-
Marquez, 2005). A prominent ridge common to all
hadrosaurines, often referred to as the quadrate but-
tress, is found on the posterior aspect of the quadrate
head (Fig. 12). The quadrate buttress present on
G. monumentensis (RAM 6797), G. incurvimanus
(TMP 80.22.1), and G. notabilis (CMN 2278) is much
larger than that described by Prieto-Marquez (2005)
for Brachylophosaurus; however, it should be noted
that the size of this feature varies substantially
amongst individuals of the same species (e.g. UMNH
VP 16667 and UMNH VP 16668). Medially, the
Figure 12. Gryposaurus monumentensis gen. et sp. rounded pterygoid wing is nearly symmetrical
nov. (RAM 6797) right quadrate: A, lateral view; B, medial
(Fig. 12).
view. mc, mandibular condyle; ptw, pterygoid wing; qb,
quadrate buttress; qh, quadrate head; qjn, quadratojugal
Pterygoid: The pterygoid, the largest element of
notch; qs, quadrate shaft. Scale bar, 5 cm.
the palate, contacts the quadrate posteriorly, the
basisphenoid medially, and the ectopterygoid,
ventrally. The basipterygoid processes diverge lat- palatine, and vomer anteriorly. In general, it is quite
eroventrally at approximately 30° from the vertical, similar to that described by Horner (1992) for Pro-
approximately equivalent to G. notabilis (ROM 873). saurolophus, and nearly identical to the pterygoid of
The transverse ridge connecting the basipterygoid G. incurvimanus (Parks, 1920). Therefore, only new
processes is not as well developed as in Brachylopho- information pertaining to the pterygoid is provided
saurus. However, a short, stout, anteroposteriorly here. The posterodorsal quadrate process is tall,
flattened prong descends vertically from the midline rising to nearly the dorsal-most surface of the quad-
of the transverse ridge (Fig. 11A), a character shared rate. This condition is present in Prosaurolophus and
with Brachylophosaurus, cf. Kritosaurus (YPM-PU apparently Edmontosaurus (Horner, 1992), but differs
16970; Horner, 1992), Edmontosaurus (Lambe, 1920; from that of G. latidens, which has a shorter dorsal
Horner, 1992), G. notabilis (ROM 873), and G. incur- quadrate process (Horner, 1992). The entire palatal
vimanus (TMP 80.22.1) (basisphenoids are unknown arch is preserved; as viewed laterally, it forms a
from G. latidens). The presence of this character is slightly raised sigmoidal contact with the palatine on
considered primitive as it is present on at least the pterygoid anterodorsal contact facet. The ptery-
Camptosaurus, Iguanodon (Norman, 1980: Fig. 5), goid contacts the vomers medial to the palatal arch.
and Bactrosaurus. The ventral prong may be absent The ventral border of the element overlaps the
in Prosaurolophus (Horner, 1992). Anteriorly, the maxilla on the large pterygoid process, producing a
parasphenoid process forms the apex of the basisphe- more substantial contact than in Prosaurolophus
noid, as in other hadrosaurids (Fig. 11B). (Horner, 1992) and Brachylophosaurus (Prieto-
Marquez, 2005).
PALATOQUADRATE COMPLEX Palatine: The palatine unites the dorsally positioned

General: RAM 6797 preserves all palatal elements, pterygoid with the maxilla ventrally through two
including the paired vomers. broad articulation facets. The arching pterygoid facet
encompasses the entire dorsal surface of the palatine,
Quadrate: The quadrate of G. monumentensis is whereas the ventral maxilla facet is long and
straight and robust (Fig. 12). The mandibular condyle straight. Posteriorly, the pterygoid facet is concave

Figure 14. Predentary of RAM 6797 in left anterolateral
view. D, dentary; nf, nutrient foramen; pdd, predentary
denticles; pdlp, predentary lateral process; pdvp, preden-
tary ventral process. Scale bar, 5 cm.
ventral lobe is excavated in RAM 6797, revealing two

Figure 13. Right vomer of Gryposaurus monumenten-
sis gen. et sp. nov. in lateral view: A, RAM 6797; B,
notches unobserved in other taxa (Fig. 13A).
UMNH VP 13970. The broken line marks the estimated
position of the dorsal margin of the anteroventral excava- MANDIBULAR COMPLEX
tion. ave, anteroventral excavation; mxa, maxillary articu- Predentary: The predentary is massive, deep, and
lation; pma, premaxillary articulation; pvl, posteroventral scoop-shaped (Fig. 14) compared with that of other
lobe. Scale bar, 5 cm. hadrosaurids. Several authors have noted the close
similarities shared in the predentaries of Kritosaurus
and Gryposaurus (Lull & Wright, 1942; Kirkland
dorsomedially, whereas it appears nearly straight in et al., 2006). Other hadrosaurines tend to have shal-
an isolated Brachylophosaurus palatine (MOR 1071- lower and more gracile predentaries, particularly
7-16-98-248-S), and is longer than illustrated for Brachylophosaurus (Prieto-Marquez, 2005), in which
P. blackfeetensis (Horner, 1992). the oral margin is straight and squared off anteriorly
instead of rounded as in Gryposaurus. The oral
Vomer: The vomer is rarely preserved in hadrosau- margin of the G. monumentensis predentary pos-
rids, making comparisons difficult. The vomer of sesses several large (width, 2 cm; height, 1.5 cm),
G. monumentensis is narrow anteriorly where it con- vertically orientated, anteroposteriorly compressed,
tacts the premaxilla and maxilla, and dorsoventrally clover-shaped denticles (Fig. 14). These denticles are
expanded posteriorly, where it meets the pterygoid. not seen on the predentary of Kritosaurus (AMNH
The laterally positioned premaxillary and maxillary 5799), other species of Gryposaurus, or in any other
articular facets are flat and occur near the anterior hadrosaurid species; therefore, they are regarded
tip of the element (Fig. 13). There seems to be little here as an apomorphy of G. monumentensis. Later-
variation in the morphology of the anterior vomer in ally, a dorsoventrally wide process extends posteri-
hadrosaurines (Lambe, 1920; Horner, 1992). Posteri- orly, overlapping the lateral side of the dentary. The
orly, the vomer remains subhorizontal on its dorsal process bears a depression on its anterior region that
edge, whereas it expands ventrally to posteroven- accepts the ventrolateral deflection of the premaxil-
trally into a lobe-shaped process (Fig. 13) that most lary oral margin. The dorsal and ventral edges of the
resembles the conformation in Brachylophosaurus predentary lateral process terminate subequally, such
(CMN 8893). The morphology of the posterior end that the longer ventral margin nearly reaches the
differs substantially from cf. Kritosaurus (YPM-PU tooth row. In ventral view, two wide flat prongs
16970; Horner, 1992) in that the dorsal margin is project posteriorly from the ventral midline of the
smooth edged, not rugose and flared as in the latter, predentary to secure the element to the predentary
and the ventral region is rounded rather than sub- shelf of the dentary (Fig. 14).
rectangular. The vomer of Maiasaura (OTM F138)
also varies from that of Gryposaurus in that both the Dentary: Overall, the dentary of the new taxon is
dorsal and ventral margins are expanded in the ver- robust and massive, yet otherwise similar to that of
tical plane. In addition, the anterior margin of the other Gryposaurus taxa (Fig. 1). It contacts the pre-
dentary anteriorly, the angular and splenial medially,

and the surangular posteriorly. In lateral view, the
dentary downturns at the anterior end of the tooth
row slightly more than in other Gryposaurus species,
and in contrast with the relatively straight dentary of
Edmontosaurus (Lambe, 1920) and Shantungosaurus
(Hu, 1973). Numerous foramina penetrate the lateral
side dentary body, including a large anterior dentary
foramen. There are over 40 tooth positions in the jaw,
with four to five replacement teeth in each alveolus.
The coronoid process is faintly inclined anteriorly.

Its dorsal margin appears more rounded than that
seen in Brachylophosaurus (Prieto-Marquez, 2005),
Prosaurolophus (Horner, 1992), or Edmontosaurus
(Lambe, 1920). Ontogenetically, the dentary appears
to have undergone substantial change, including a
major increase in robustness and an alteration of the
dorsal surface of the diastema (i.e. the contact surface
for the predentary), from dorsally concave in juveniles
and subadults (i.e. UMNH VP 13970) to dorsally
convex in adult specimens (i.e. RAM 6797). Figure 15. Right surangular of Gryposaurus monu-
mentensis gen. et sp. nov. (RAM 6797): A, lateral view;
Surangular: Five elements contact the surangular: B, dorsal view. ap, articular process; asn, anterior suran-
the dentary, splenial, angular, articular, and quad- gular notch; mc, medial concavity; mr, median ridge; msp,
rate. The articular adheres to the posterior border of median surangular process; qas, quadrate articular
the surangular. The splenial and angular are posi- surface; scp, surangular coronoid process. Scale bar, 5 cm.
tioned dorsally and ventrally, respectively, about a
small sagittal ridge along the medial side of the
surangular. Laterally, the mandibular condyle of the orly, and the jugal dorsally. In other species of
quadrate rests in a dorsally orientated depression Gryposaurus, the foramen is subcircular, whereas, in
(Fig. 15). A midline sagittal ridge separates the G. monumentensis, it is ovoid (Fig. 5; see below).
medial and lateral halves of the surangular from each
other (Fig. 15B). Midlength along this ridge, an
POSTCRANIAL SKELETON
anteroposteriorly broad process ascends anterodor-
sally (Fig. 15A). The morphology of this process in The postcranial skeleton referred to G. monumenten-
G. monumentensis differs from the more triangular sis, UMNH VP 12265, is largely unprepared and will
process seen in Prosaurolophus (Horner, 1992), not be described here. In general, the skeleton is
Brachylophosaurus (Prieto-Marquez, 2005), and much more robust than that seen in G. incurvimanus
Edmontosaurus (Lambe, 1920), in that the posterior (Parks, 1920) and Brachylophosaurus (Prieto-
region is dorsoventrally reduced and the anterodorsal Marquez, 2007). The type specimen of G. notabilis is
corner is flared dorsally to form a small flange. The the only specimen of this species with a postcranial
anterior region of the surangular is shaped laterally skeleton, but, unfortunately, offers little comparative
by a dorsally projecting, mediolaterally compressed, material because it is not well preserved.
surangular coronoid process. This process is shorter
and more robust than that in Brachylophosaurus
PHYLOGENETIC ANALYSIS
(Prieto-Marquez, 2005). A small notch, here termed
the anterior surangular notch, is present on the A phylogenetic analysis was conducted in order to
mediolateral margin of the element (Fig. 15B). This assess the relationships of G. monumentensis within
notch is absent in Brachylophosaurus and P. black- Hadrosaurinae. The matrix for this analysis was
feetensis (Horner, 1992), but unobservable and unveri- based on that presented by Horner, Weishampel &
fiable on any other specimen of Gryposaurus because Forster (2004), supplemented with characters from
of contact with the dentary. However, several basal Weishampel et al. (1993) and novel characters result-
iguanodontians possess a second foramen in the same ing from this study (Appendix 1). A total of 120 char-
location on the surangular (Norman, 1998). acters was analysed amongst the 15 taxa using a
The mandibular foramen is composed of the suran- branch and bound search under ACCTRAN optimiza-
gular anteriorly and ventrally, the quadrate posteri- tion in PAUP version 4.0b10 (Swofford, 2002). Tree

Figure 16. Strict consensus of the seven most parsimonious phylogenetic trees demonstrating the relationship of
Gryposaurus to other hadrosaurines. A total of 15 taxa and 120 characters was analysed in MacClade version 4.06
(Maddison & Maddison, 2000) and PAUP version 4.0b10 (Swofford, 2002) using the ACCTRAN optimization. Iguanodon
and Eolambia were considered as outgroups. Letters on branches denote clades with the character support listed below,
and numbers following branch designations refer to the bootstrap values (1000 replicates) associated with that branch.
Branch A is supported by characters 1, 29, 35, 37, 59, 64, 76, and 86 (bold, unambiguous characters). Branch B is
supported by characters 24, 43, 53, 65, 66, 67, and 79. Branch C is supported by characters 29, 40, 54, and 62. Branch
D is supported by characters 12, 14, 31, 42, and 65. Branch E is supported by characters 10, 22, 23, 40, 45, 58, 62, 88,
and 100. See Appendix 1 for character descriptions and Appendix 2 for character codings.
manipulation was performed and the character state primitive Gryposaurus taxon based on the large size
distribution was analysed using MacClade version of the dentary teeth. Numerous other authors
4.06 (Maddison & Maddison, 2000). (Weishampel et al., 1993; Horner et al., 2004; Prieto-
Figure 16 shows the strict consensus tree derived Marquez et al., 2006) have also considered this trait
from the seven most parsimonious trees. The tree as primitive. Horner (1992) further explains that
posits two main clades of hadrosaurine hadrosaurids G. latidens possesses traits of both G. incurvimanus
– one clade consisting of Saurolophus + Prosauro- and G. notabilis, which makes refinement of the
lophus + Edmontosaurus and the other comprising phylogenetic tree more problematic. Gryposaurus
Naashoibitosaurus + Gryposaurus + Maiasaura + Bra- monumentensis, on the other hand, is plausibly
chylophosaurus. Lophorothon rests at the base of the more closely related to G. notabilis than to the other
Hadrosaurinae clade (Fig. 16). Gryposaurus falls out species. The two taxa share similar general skull
within a clade including Maiasaura + Brachylopho- morphology, both are more robust than either
saurus, supported by four ambiguous characters G. latidens or G. incurvimanus, and both possess the
(Fig. 16). Character support for this clade is weak following features: increased size of the premaxillary
owing to a lack of phylogenetic resolution within lip; sigmoidal maxillary lateroventral margin; and
Hadrosaurinae. Therefore, more data are needed in pronounced jugal tubercle. Yet, all species of this
order to support or refute this grouping. All four genus are quite comparable, limiting the resolution
species of Gryposaurus form a polytomy supported of specific relationships.
by five ambiguous characters (Fig. 16). Horner The overall tree topology presented here differs in
(1992) hypothesized that G. latidens is the most several aspects from the analyses of Horner et al.
(2004) and Prieto-Marquez et al. (2006). In the former 5799) and those taxa that have been referred to
analysis, Naashoibitosaurus and Saurolophus pair this genus [Naashoibitosaurus (sensu Horner, 1992;
together within a small clade, whereas Prosaurolo- Horner et al., 2004) and Anasazisaurus (sensu
phus and Edmontosaurus are posited in a larger clade Horner, 1992)] is required before the taxonomy and
that includes Brachylophosaurus and Gryposaurus. phylogenetic relationships of these hadrosaurids can
Lophorothon remains basal to the clade containing be assessed. Nevertheless, following the synonymy of
Gryposaurus in Horner et al. (2004), whereas it is Horner (1992) and Horner et al. (2004), we currently
positioned here at the base of Hadrosaurinae. In this find a clear distinction between Gryposaurus and
study, Gryposaurus is closely aligned with Brachylo- Kritosaurus based on: (1) the distinct morphology of
phosaurus and Maiasaura, by contrast with the the maxilla in Anasazisaurus (Lucas et al., 2006); (2)
results of Horner et al. (2004) and Prieto-Marquez differing morphology of the nasal ornamentation –

et al. (2006), which both suggest a much closer rela- which is a posteriorly folded, rugose, triangular crest
tionship between Gryposaurus and Prosaurolophus. in Anasazisaurus, and a raised rugose region in
Naashoibitosaurus (Horner, 1992; Hunt & Lucas,
1993; Lucas et al., 2006); (3) a circumnarial depres-
sion extending to near the orbits in Anasazisaurus,
DISCUSSION
much further than that observed in Gryposaurus
TAXONOMIC AND EVOLUTIONARY IMPLICATIONS (Horner, 1992; Lucas et al., 2006; this paper); (4)
With the addition of G. monumentensis, Gryposaurus different nasofrontal sutures – the frontals insert a
now represents the most taxonomically diverse had- small process into the nasal in Naashoibitosaurus
rosaurid genus known. A revised generic diagnosis instead of the opposite condition in Gryposaurus
unites the four species by a suite of synapomor- (Horner, 1992); and (5) opposing shapes of the ischia
phies, including: (1) dorsolateral flare on the medial – the shaft is straight in Gryposaurus, whereas
side of the premaxillary lateral process (Figs 3, 5); referred specimens of Kritosaurus have dorsally
(2) mediolaterally compressed, arched nasal orna- curved shafts (Kirkland et al., 2006).
mentation; and (3) sigmoidal frontal–nasal suture in Table 1 presents the ratio of the average orbital
which the nasals insert a small process into the area to the infratemporal fenestra area for the three
frontals (Fig. 6). Previous generic diagnoses of Gry- species of Gryposaurus. This reveals that G. monu-
posaurus included only a single unique feature – the mentensis is the only taxon whose ratio is greater
nasal arch – together with several other features than one (1.13); that is, the orbit is significantly
(e.g. dorsoventrally deep skull and infratemporal larger than the infratemporal fenestra, further con-
fenestra larger than the orbit) that pertain to some firming that this character is not a genus-level syna-
other hadrosaurine taxa (i.e. Kritosaurus; Lull & pomorphy. Gryposaurus sp. (UMNH VP 16667) has a
Wright, 1942). subequal orbit to infratemporal fenestra area ratio of
Gryposaurus monumentensis exemplifies the inad- 0.97, whereas the two G. notabilis specimens and
equacy of using the infratemporal fenestra in generic both G. incurvimanus specimens sampled had a
diagnoses, as this feature is significantly smaller than maximum value of 0.70, indicating that the orbit is
the orbit in this taxon (see below). Horner (1992) also significantly smaller than the infratemporal fenestra.
addressed the morphology and use of the infratempo- The fact that G. notabilis and G. incurvimanus both
ral fenestra in the generic diagnoses of Gryposaurus possess an orbit to infratemporal fenestra area ratio
and Kritosaurus. He recognized subtle differences of 0.70 weakens the hypothesis that these two taxa
between the taxa, determined that the infratemporal are part of an ontogenetic series instead of distinct
fenestra was not a taxonomically informative charac- taxa. If they were part of a series, it would be
ter of these hadrosaurines, and ultimately eliminated expected that the infratemporal fenestra would be
it from the diagnoses of both genera. smaller relative to the orbit in G. notabilis than in
With regard to the taxonomic relationship of Gry- G. incurvimanus.
posaurus to Kritosaurus – that is, whether the two The decrease in the size of the infratemporal fenes-
taxa are synonymous, sister taxa, or convergent in tra in G. monumentensis appears to have been asso-
morphology – the present analysis lends additional ciated with a substantial restructuring of the
evidence, but cannot fully resolve this issue, because elements in the posterior region of the skull. Mini-
of morphological uncertainties within Kritosaurus. mally, this transformation involved shortening of the
The two genera are superficially similar in possessing postorbital and squamosal processes that comprise
deep skulls with a massive predentary and (for most the supratemporal bar and rotation of the jugal pos-
Gryposaurus species) infratemporal fenestrae that terior process dorsally. In addition, other probable
are larger than the orbit. However, detailed anatomi- consequences of this restructuring include slight
cal study of the type specimen of Kritosaurus (AMNH anteroposterior shortening of the jugal posterior
process, anteroposterior shortening of the parietal

Table 1. List of skull measurements for Gryposaurus monumentensis, G. incurvimanus, and G. notabilis. Skull depth at the nasal was measured vertically
measurements. Average orbital and ITF areas were obtained by averaging the area from both sets of measurements. The ITF and orbital ratios were calculated
the subsequent measurement taken perpendicular to the initial measurement. In addition, length and width measurements were taken at 45° to the previous
from the dorsal-most surface of the nasal to the base of the dentary. Orbital dimensions were obtained from measuring the longest vertical length and longest
width perpendicular to the length. The first infratemporal fenestra (ITF) length was taken along the longest diagonal length of the fenestra, not vertically, with
UMNH VP 16667
Gryposaurus sp.
(this conformation is inferred, as the parietal of
G. monumentensis is currently not known), and par-
allel shortening of the surangular and/or dentary
0.59
0.62
0.92
0.53
0.97
relative to the skull depth (Table 1). The latter modi-
90.0
53.3
55.9
425.2
439.3
fication appears to have been necessary, as decreasing
the supratemporal bar decreases the total skull
length and thereby necessitates an equivalent
decrease in the length of the mandibular complex.
CMN 2278
Observation of the mandibular foramen in other

species of Gryposaurus reveals a subcircular opening
0.61
0.65
0.67
0.55
0.67
80.0
48.5
52.0
246.9
326.9

that contrasts with the oval foramen present in
G. monumentensis (RAM 6797; Fig. 5). Anteroposte-
rior shortening of the surangular, not the dentary,
would be necessary in order to achieve the shape
G. notabilis
change from a subcircular to an oval mandibular

ROM 873
foramen.
0.60
0.68
0.83
0.67
0.70
75.0
45.0
51.0
290.3
416.0
Overall, the general skull morphology of G. monu-

mentensis most closely resembles that of G. notabilis,
although the skull of the former taxon appears boxier
than the latter. The skull depth to length ratios
presented in Table 1 confirm that the skull shape of
ROM 764
G. monumentensis is in fact boxier than that of G. no-

0.84
0.57
0.68
40.0
188.0
275.5
tabilis – that is, the ratios are closer to one in the new
–
–
–
–
Utah taxon. However, this ratio is less than that

exhibited by G. incurvimanus, which possesses a qua-
drangular skull of nearly subequal maximum skull
G. incurvimanus
depth to length ratio of 0.94 (Table 1).

RTMP 80.22.1
Gryposaurus monumentensis can be further distin-

guished from other species of the genus by the more
0.82
0.94
0.86
0.77
0.70
robust nature of the skull and lower jaws, a feature

42.5
35.0
40.0
166.2
215.5
that is perhaps best revealed in the predentary,

dentary, and premaxilla (Figs 1, 3). Most of these
unique features of the skull and lower jaws of
G. monumentensis are reasonably inferred as adapta-
G. monumentensis
tions for generating and absorbing heightened bite

forces. These features include: (1) a more robust
dentary, predentary, and premaxilla; (2) a steeper
RAM 6797
by (smallest measure/largest measure) in each set
angled premaxillae and shorter skull length relative

to depth, both of which would have permitted a more
0.65
0.72
0.76
0.37
1.13
76.0
49.5
54.6
342.3
301.9
vertical bite action and increased the bite force of the

anterior region of the skull; (3) increased rugosity on
the midline of the premaxilla to more securely attach
the upper rhamphotheca; and (4) large, pronged den-
ticles of the predentary, plausibly serving as an
Skull depth at nasal (NSD) (cm)
increased attachment area for the lower rhamphoth-

Max. skull length (MSL) (cm)
Max. skull depth (MSD) (cm)
eca. Whether or not these characters indicate a

Av. orbit area (AOA) (cm2)
Av. ITF area (ITFA) (cm2)
tougher, more fibrous diet relative to that of closely

related species cannot be ascertained from this
strictly morphological assessment. However, the evi-
dence is intriguing and might profitably be the focus
Av. orbit ratio
of a future functional analysis. Nonetheless, given the

Av. ITF ratio
AOA/ITFA
MSD/MSL
distinction in predentary shape between Gryposaurus

NSD/MSL
species and other hadrosaurid taxa, members of this

genus may have been more capable of processing a
wider variety of plant material.
ONTOGENETIC CONSIDERATIONS the subject of a separate study, more closely

Although G. monumentensis occurs later in time and resemble those of G. notabilis. The presence of two,
in a different location than G. notabilis and G. in- plausibly stratigraphically separated species of Gry-
curvimanus, the possibility exists that the speci- posaurus in the Kaiparowits Formation (although
mens currently at hand comprise an ontogenetic more specimens are needed to determine how much
series of one species, as opposed to distinct species. stratigraphic overlap, if any, is present between the
The only line of evidence supporting the notion that two taxa) is significant, in that it is strongly sug-
these taxa are part of a growth series beginning gestive of within-lineage faunal turnover from a
with G. incurvimanus and ending with G. monumen- single geological formation. The turnover proposed
tensis is that overall skull gracility increases from in the Kaiparowits is similar to patterns observed in
G. incurvimanus to G. monumentensis. However, dinosaurian faunas in northern Western Interior

several other pieces of evidence conflict with the Basin strata, such as the Two Medicine Formation
ontogeny hypothesis. Hadrosaurids grow continu- (Horner, Varricchio & Goodwin, 1992) and the
ously throughout their life (Horner, Ricqles & Dinosaur Park Formation (Ryan & Evans, 2005;
Padian, 2000); therefore, if G. monumentensis is the Evans et al., 2006). This discovery indicates that
culmination of the ontogenetic series, the maximum approximately 75 Mya, one species of Gryposaurus
skull length should be longer than that observed in inhabiting southern Utah was replaced by G. monu-
G. notabilis. Table 1 shows that the type specimen mentensis. Presently, there is no evidence that this
of G. notabilis (CMN 2278) is larger than the type of replacement correlates with changing environmental
G. monumentensis (RAM 6797), and yet retains the conditions, and clearly not enough of the fossil
characteristics seen on a slightly smaller G. notabi- record of these taxa is known regionally to differ-
lis specimen (ROM 873, which is 1 cm shorter than entiate between other patterns of replacement, such
RAM 6797). Further, the infratemporal fenestra as habitat tracking or extinction.
does not slowly decrease in size throughout the Hadrosaurine material collected from the Lower
hypothetical growth series, as discussed above. It Shale Member of the Aguja Formation in southern
should be assumed that, during ontogeny, the Texas may be attributable to yet another new species
infratemporal fenestra would not remain the same of Gryposaurus (Kritosaurus of Wagner, 2001; Wagner
size for the majority of the hadrosaurid’s life and & Lehman, 2001). This material possesses the char-
then dramatically decrease in size (including modi- acteristic Gryposaurus arched nasal ornamentation,
fication of the entire posterior skull). It seems more together with other features that are cited by Wagner
parsimonious to assume that the difference in size (2001) as evidence for a close relationship to G. lati-
and posterior skull structure seen in G. monument- dens. In addition, Wagner (2001) describes maxillae
ensis is the result of speciation, not ontogeny. from the Upper Shale Member of the Aguja that
Finally, subadult specimens of G. monumentensis closely resemble Gryposaurus (Kritosaurus of Wagner,
(UMNH VP 13970) that coincide in size with those 2001) notabilis.
of G. incurvimanus (TMP 80.22.1) demonstrate that The overall biostratigraphic pattern of Gryposaurus
the nasal morphology differs between the two taxa. reveals that the genus persisted for at least five
More specifically, the nasal arch in the subadult million years, significantly longer than any other
G. monumentensis (UMNH VP 13970) is compara- taxon within Hadrosauridae, with the exception of
tively larger and positioned more posteriorly than Edmontosaurus. Gryposaurus latidens first occurs in
that in G. incurvimanus, more similar to the nasal the lower Two Medicine Formation of Montana (c.
seen in adult specimens of G. notabilis and 80 Mya), approximately four million years prior to the
G. monumentensis. In short, morphological evidence appearance of the other species of Gryposaurus [but it
indicates that G. monumentensis is probably a new also occurs throughout the Two Medicine Formation
taxon and not a data point along the ontogenetic (Varricchio, 1995) at the same time as other Grypo-
growth of one Gryposaurus species. saurus species]. Ryan & Evans (2005) documented the
co-occurrence of G. notabilis and G. incurvimanus in
the lower third of the Dinosaur Park Formation in
BIOGEOGRAPHY AND BIOSTRATIGRAPHY Alberta, Canada, at approximately 76 Mya. The speci-
Fossils of G. monumentensis have been found within mens of Gryposaurus sp. found in the lower portion of
the middle unit of the Kaiparowits Formation. Had- the Kaiparowits Formation occur equivalently in time
rosaurine materials recovered from near the base of with G. notabilis and G. incurvimanus in Dinosaur
the formation and a portion of the lower middle Park, whereas G. monumentensis occurs in somewhat
unit, including an articulated skull (UMNH VP younger sediments dating between 75 and 74 Mya,
16667), clearly pertain to Gryposaurus, but not to apparently coeval with P. maximus in the Dinosaur
G. monumentensis. These materials, which will be Park Formation (Gates & Evans, 2005).
CONCLUSIONS geographical range (exceeding 2000 km north to

south) is one of the largest amongst late Campanian
Gryposaurus monumentensis, recovered from the dinosaur genera.
middle portion of the Kaiparowits Formation, repre-
sents a new hadrosaurine from the late Campanian of
southern Utah. It is the fourth Gryposaurus species to ACKNOWLEDGEMENTS
be named, making this genus the most diverse within
We express sincere thanks to the following: Duncan
Hadrosauridae. Previous attempts at diagnosing Gry-
Everhart, who discovered the holotype specimen of
posaurus relied heavily on such characters as the
G. monumentensis; the students, teachers, and volun-
distinguishing nasal arch, the shape of the frontal
teers of the ALF Museum Peccary Society, who exca-
nasal contact, and a larger infratemporal fenestra
vated the holotype specimen; Don Lofgren for support

relative to the orbit. The latter character is absent
and unrestricted study of the holotype specimen; Alan
in G. monumentensis, which possesses a narrow
Titus for his logistical support, enthusiasm, and keen
infratemporal fenestra, nearly half the size of the
eye for discovering new specimens; Grand Staircase-
orbit; thus, this discovery required amendment of
Escalante National Monument and the Bureau of
the generic diagnosis of Gryposaurus to exclude this
Land Management for permits and funding; Mike
feature. In addition, a flaring on the medial side of the
Getty and the UMNH laboratory volunteers and field
premaxillary lateral process was found to be a diag-
crew for excavating the hadrosaurid material from
nostic feature of Gryposaurus. Gryposaurus monu-
GSENM; Eric Lund, Sharon Walkington, and the
mentensis is distinguished from the other species of
volunteers of UMNH for their devoted preparation
Gryposaurus by the possession of a boxy (i.e. rela-
work; David Evans, Albert Prieto-Marquez, and Jack
tively short length to height ratio), more robust skull
Horner for enlightening hadrosaurid conversations;
and lower jaws, a relatively small infratemporal
Lindsay Zanno for ongoing support and illustrations
fenestra, the presence of a median flange on the
that greatly improved the manuscript; and Jim
surangular, and large prongs on the oral margin of
Gardner, Jack Horner, Jim Kirkland, Albert Prieto-
the predentary. Most of the unique morphologies of
Marquez, Kevin Seymour, and Kieran Shepherd for
G. monumentensis appear to be related not to classi-
access to specimens in collections and photographs.
cally appointed display structures, but rather to
Reviews by Tony Ekdale, Jack Horner, and two
extreme overall robustness of the skull (producing a
anonymous reviewers greatly improved the quality of
more dramatic difference between congeners than
the manuscript. Additional funding was provided by
observed in most other hadrosaurid taxa), plausibly
the Discovery Channel and the Jurassic Foundation.
related to feeding adaptations.
A phylogenetic analysis of Hadrosaurinae that
includes the new taxon resulted in Gryposaurus
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Tech University.
four or five teeth (0); long, equal to approximately
Wagner JR, Lehman T. 2001. A new species of Kritosaurus one-fifth to one-quarter of length of tooth row (1);
from the Cretaceous of Big Bend National Park, Brewster extremely long, equal to approximately one-third of
County, Texas. Journal of Vertebrate Paleontology 21 tooth row (2).
(Suppl.): 110A–111A. 11. Dentary tooth row, posterior extent of tooth row
Weishampel DB, Jenson JA. 1979. Parasaurolophus (Rep- relative to apex of coronoid process: tooth row termi-
tilia: Hadrosauridae) from Utah. Journal of Paleontology nates even with or anterior to apex (0); tooth row
53: 1422–1427. terminates posterior to apex (1).
Weishampel DB, Norman DB, Grigorescu D. 1993. Tel- 12. Dentary, orientation of dentary anterior to tooth
matosaurus transsylvanicus from the Late Cretaceous of row: moderately downturned, dorsal margin of pre-
Romania: the most basal hadrosaurid dinosaur. Palaeontol- dentary rests above ventral margin of dentary body
ogy 36: 361–385. (0); strikingly downturned, dorsal margin of anterior
Zanno LE, Sampson SD. 2005. A new oviraptorosaur dentary extends below the ventral margin of dentary
(Theropoda, Maniraptora) from the Late Cretaceous (Cam- body, premaxillary bill margin extends well below
panian) of Utah. Journal of Vertebrate Paleontology 25: level of maxillary tooth row (1). (Adapted from Horner
897–904. et al., 2004: character 11.)
13. Dentary tooth row, shape in occlusal view: bowed
lingually, curves in towards coronoid process (0);
straight (1).
APPENDIX 1
14. Predentary shape: deep and robust (0); gracile
The character descriptions listed below are those and shovel-shaped (1). (Adapted from Horner et al.,
presented by Horner et al. (2004), unless otherwise 2004: character 13.)
15. Predentary shape: arcuate anterior margin, neu- 27. Premaxillae, oral margin with a ‘double layer’
rovascular foramina large and located near midline morphology consisting of an external denticle-
of predentary body, dorsally directed spike-like den- bearing layer seen externally and an internal
ticles on anterior margin that fit into slots on palatal layer of thickened bone set back slightly
underside of premaxilla (0); straight to gently from the oral margin and separated from the den-
rounded anterior margin, numerous nutrient ticulate layer by a deep sulcus bearing vascular
foramina across entire anterior margin, loss of foramina: absent (0); present (1).
spike-like denticles and gain of rounded, triangular 28. Premaxilla, outer (accessory) narial fossa ante-
denticles that project anteriorly and fit into a con- rior to circumnarial fossa: absent (0); present, sepa-
tinuous transverse slot on underside of premaxilla rated from circumnarial fossa by a strong ridge (1).
(1). 29. Premaxilla-nasal: shape between anterior-most

16. Predentary triturating surface, orientation: hori- point of premaxilla and posterior region of external
zontal, oral margin of premaxilla rests on dorsal nares: slightly arched (0); highly arched (1); slightly
predentary (0); canted dorsolaterally to form a concave to horizontal (2) (new).
nearly vertical surface, oral margin of premaxilla 30. Premaxillary posterior processes (PM1, PM2)
broadly overlaps lateral surface of predentary (1). and construction of nasal passages: posterodorsal
17. Angular size: large, deep, exposed in lateral premaxillary process short, posterodorsal and pos-
view below the surangular (0); small, dorsoventrally teroventral processes do not meet posterior to exter-
narrow, exposed only in medial view (1). nal nares, nasal passages not enclosed ventrally,
18. Coronoid bone: present (0); absent (1). anterior nasal passage roofed by the nasal, external
19. Coronoid process configuration: apex only nares exposed in lateral view (0); posteroventral and
slightly expanded anteriorly, surangular large and posterodorsal processes elongate and join behind
forms much of posterior margin of coronoid process external opening of narial passages to exclude
(0); dentary forms nearly all of greatly anteroposte- nasals, nasal passages completely enclosed by
riorly expanded apex, surangular reduced to thin tubular premaxillae, left nasal passage divided from
sliver along posterior margin and does not reach to right passage, external nares not exposed in lateral
the distal end of the coronoid process (1). view (1).
20. Posterior extent of the posteroventral dentary: 31. Premaxilla, lateral process flares dorsally into
ends even with or anterior to apex of coronoid external naris: absent (0); present (1) (new).
process (0); posteriorly expanded to terminate well 32. External nares length to basal skull length
behind the coronoid process (1). ratio: 20% or less (0); 30% or more (1).
21. Surangular foramen: present (0); absent (1). 33. External nares, composition of posterior-most
22. Premaxilla, width at oral margin: narrow, apex: formed entirely by nasal (0); formed equally
expanded laterally to less than twice width at nar- by nasal (dorsally) and premaxilla (ventrally) (1);
rowest point (postoral constriction), margin orien- formed only by premaxilla (2). (Adapted from
tated nearly vertically (0); expanded transversely to Horner et al., 2004: character 29.)
more than twice postoral width but not more than 34. External naris: posterior ventral margin com-
interorbital width, margin flared laterally into a posed of premaxilla (0); posterior margin composed
more horizontal orientation (1); further expanded of nasal up to 25% of total naris length (1) (new).
transversely to width subequal to that across jugal 35. External naris: anterodorsal corner composed of
arches (2). only premaxilla dorsal process (0); composed of both
23. Premaxilla, undercut (‘reflected’) rim around premaxilla dorsal process and nasal anterior process
oral margin: absent (0); present (1). (1); entire naris formed by premaxilla (2) (new).
24. Premaxillary anterior bill margin shape: 36. Circumnarial fossa, posterior margin: absent (0);
horseshoe-shaped, forms a continuous semicircle present (1).
that curves smoothly to postoral constriction (0); 37. Circumnarial fossa, posterior margin morphol-
broadly arcuate across anterior margin, constricts ogy: absent (0); present, lightly incised into nasals
abruptly behind the oral margin (1). and premaxilla, often poorly demarcated (1); pre-
25. Premaxillary foramen ventral to anterior sent, well demarcated, deeply incised, and usually
margin of external nares that opens onto the palate: invaginated (2).
absent (0); present (1). 38. Nasals and anterodorsal premaxilla in adults:
26. Premaxilla, accessory foramen entering premax- flat, restricted to area anterior to braincase, cavum
illa in outer narial fossa, located anterior to pre- nasi small (0); premaxilla extended posteriorly and
maxillary foramen: absent (0); present, empties into nasals retracted posteriorly to lie over braincase in
common chamber with premaxillary foramen, then adults resulting in a convoluted, complex narial
onto the palate (1). passage and hollow crest, cavum nasi enlarged (1).
39. Solid nasal crest over snout or braincase (does orbital rim (0); expanded dorsoventrally in front of
not house a portion of the nasal passage): absent orbit, lacrimal pushed dorsally to lie completely above
(0); present (1). the level of the maxilla, jugal forms lower portion of
40. Solid nasal crest, association with posterior orbital rim (1).
margin of circumnarial fossa: absent (0); solid crest 57. Jugal, shape of anterior end: with distinct anteri-
present but circumnarial fossa does not excavate orly pointed process fitting between the maxilla and
side of crest, fossa terminates anterior to solid crest lacrimal (0); point truncated, smoothly rounded ante-
(1); solid crest present, excavated laterally by cir- rior margin (1).
cumnarial fossa (2). 58. Jugal, anteriorly pointed process: absent (0);
41. Solid nasal crest, composition: absent (0); solid present, process restricted to dorsal portion of jugal,
crest present, composed of nasals (1). (Adapted from anterior jugal appears asymmetrical (1); present,

Horner et al., 2004: character 40.) process centred on anterior jugal, anterior jugal
42. Solid nasal crest, creating a mediolaterally com- appears symmetrically triangular in shape (2).
pressed arch anterior to orbits: absent (0); present 59. Jugal, ventral margin of anterior process: straight
(1) (new). to slightly curved (0); sigmoidal (1); convex (2) (new).
43. External nares, shape of posterior margin: 60. Jugal, postorbital process at or nearly at 90° to the
lunate (0); ‘V’-shaped (1). body of jugal (0); highly inclined posteriorly (1) (new).
44. Maxilla, anterodorsal process: has a separate 61. Jugal, development of posteroventral flange:
anterior process that extends medial to the poster- absent, jugal expands gradually below infratemporal
oventral process of premaxilla to form part of fenestra to meet the quadratojugal–quadrate (0);
medial floor of external naris (0); anterior process present, jugal dorsoventrally constricted beneath
absent, anterodorsal margin of maxilla forms a infratemporal fenestra to set off flange anterior to
sloping shelf that underlies the premaxilla (1). constriction (1).
45. Maxilla, medial anterodorsal process: not seen 62. Jugal flange size, ratio of depth of jugal at con-
in lateral view of external nares (0); projects into striction below infratemporal fenestra to length of free
external nares in lateral view (1) (new). ventral flange on jugal: small, 0.70–0.90 (0); promi-
46. Antorbital fenestra, external opening: present nent, well offset from body of jugal, 0.55–0.66 (1).
(0); absent (1). 63. Shallow posterior jugal process: relatively deep
47. Maxillary foramen, location: opens on anterolat- posterior region of jugal posterior process (0); rela-
eral body of maxilla, exposed in lateral view (0); tively shallow posterior process (1) (Weishampel et al.,
opens on dorsal maxilla along maxilla–premaxilla 1993: character 17).
suture (1). 64. Scalloped ventral margin of posterior process of
48. Maxilla–lacrimal contact: present (0); lost or jugal: posterior process straight to slightly convex (0);
covered as a result of jugal–premaxilla contact (1). posterior process distinctly concave (1) (Weishampel
49. Maxilla–jugal contact: restricted to finger-like et al., 1993: character 18).
jugal process on posterior margin of maxilla (0); 65. Frontonasal suture at midline of skull: no distinct
jugal process of maxilla reduced to a short projec- processes (0); nasals insert small process into frontals
tion but retaining a distinct facet (1); jugal process (1); frontals insert small process between nasals (2);
of maxilla lost, anterior jugal has an extensive ver- frontals rise posterodorsally to support nasal (3) (new).
tical contact with maxilla anterior to orbit (2). 66. Frontal at orbit margin: forms part of margin (0);
50. Maxilla, location of apex in lateral exposure: well excluded by prefrontal–postorbital contact (1).
posterior to centre (0); at or anterior to centre (1). 67. Frontals, upward doming over braincase in adults:
51. Maxilla, shape of apex in lateral exposure: low and absent (0); present (1).
gently rounded (0); tall and sharply peaked (1). 68. Supraorbital articulation: freely articulate on
52. Maxilla, shape of ‘lateroventral margin’: straight orbit rim (0); fused to orbit rim or absent (1).
to nearly straight (0); sigmoidal (1); arched (2) (new). 69. Quadrates, shape of mandibular condyle: medio-
53. Prefrontal shape at anterodorsal orbit rim: pre- laterally broad, lateral and medial condyles subequal
frontal lies flush with surrounding elements (0); pre- in size (0); lateral condyle expanded anteroposteriorly
frontal flares dorsolaterally to form a thin, everted, so that condyles appear subtriangular in distal view,
wing-like rim around anterodorsal orbit margin (1). lateral condyle longer than medial one (1).
54. Prefrontal, shape: smoothly curved laterally (0); 70. Paraquadratic foramen, present (0); absent (1)
anteriorly broad with square anteromedial corner (1). (Weishampel et al., 1993: character 21).
55. Ectopterygoid–jugal contact: present (0); absent, 71. Quadratojugal notch: ventral margin of notch
palatine–jugal contact enhanced (1). extends dorsally to form an acute and well-defined
56. Jugal, expansion of anterior end below lacrimal: opening (0); well-defined notch absent, reduced to a
dorsoventrally narrow, forms little of the anterior poorly defined embayment of quadrate (1).
72. Paroccipital process and accompanying squamo- 88. Cervical centrum axial length: long (0); shortened
sal, orientation: straight and ventrally directed (0); so that axial length of centrum is less than height of
curved anteriorly (1). neural arch (1).
73. Squamosals on skull roof, separation: widely sepa- 89. Cervicals, shape of zygapophyseal peduncles on
rated (0); squamosals approach midline, separated by arches: low (0); elevated, extend well above the level of
narrow band of parietal (1); squamosals have broad the neural canal, zygapophyses long and dorsally
contact with each other (2). arched (1).
74. Squamosal, shape of posteroventral surface: shal- 90. Sacral vertebrae, number: seven or less (0); eight
lowly exposed in posterior view (0); forms a deep, or more (1).
near-vertical, well-exposed face in posterior view (1). 91. Dorsal (posterior) and sacral neural spines: short,
75. Supraoccipital, inclination: posterior surface less than three times centrum height (0); elongate,

nearly vertical (0); posterior surface inclined steeply more than three times centrum height (1).
forward at approximately 45° (1). 92. Coracoid size: large, coracoid to scapula length
76. Supraoccipital, ventral margin: bowed or ex- more than 0.2, length of articular surface greater than
panded ventrally along midline (0); horizontal, strong length of glenoid (0); coracoid reduced in length
ridge developed along supraoccipital–exoccipital relative to scapula, glenoid longer than articulation
suture (1). (1).
77. Supraoccipital–exoccipital contact: straight 93. Coracoid, shape of cranial margin: straight or
suture that meets squamosal (0); ventrolateral corner convex, biceps tubercle small (0); concave, large, later-
of supraoccipital inset into exoccipital so that supraoc- ally projecting biceps tubercle (1).
cipital is ‘locked’ between exoccipitals (1). 94. Coracoid, cranioventral process: short (0); long,
78. Basisphenoid, transverse ridge: present between extends well below the glenoid (1).
basipterygoid processes (0); absent (1) (new). 95. Scapula, shape of proximal end: dorsoventrally
79. Basisphenoid, ventral process on transverse ridge: deep, acromion process directed dorsally, articulation
present, descends from transverse ridge, ventrally or extensive (0); dorsoventrally narrow (no wider than
anteriorly inclined (0); absent (1) (new). distal scapula), acromion process projects horizontally,
80. Cranial nerve V: composed of pro-otic, laterosphe- cranioventral corner notched, articulation restricted
noid, and basisphenoid (0); composed of only pro-otic (1).
and laterosphenoid (1) (new). 96. Scapula, orientation of borders of distal blade:
81. Transverse width of the cranium in the postorbital divergent (0); subparallel to one another (1).
region in dorsal view: broad, width maintained from 97. Scapula, shape of distal end: asymmetrical, either
orbit to quadrate head (0); distinctly narrowed at dorsal or ventral border longer than the other (0);
quadrate heads (1). symmetrical, dorsal and ventral border terminate at
82. Occiput shape in posterior view: square (0); trian- same point (1).
gular, narrow dorsally, distal quadrates splay dis- 98. Deltopectoral crest: short, much less than half the
tinctly laterally (1). length of the humerus, narrows noticeably distally (0);
83. Parietal, midline ridge: straight to slightly down- extends at least to midshaft or longer, distally broad
warped along length (0); strongly downwarped, dorsal (1).
margin bends below the level of the postorbital– 99. Humeral distal condyles: mediolaterally broad,
squamosal bar (1). flare moderately from shaft of humerus (0); com-
84. Parietal crest, length: long, posterior parietal pressed mediolaterally, flare little from shaft of
narrows quickly to form the crest, crest more than half humerus (1).
the length of the supratemporal fenestrae (0); short, 100. Antebrachium length: humerus subequal to or
parietal crest narrows gradually posteriorly, crest less longer than radius (0); radius longer than humerus
than half the length of the supratemporal fenestrae (1).
(1). 101. Carpus: all elements present (ulnare, radiale,
85. Infratemporal fenestra, acute angle between intermedium, distal carpals) and fused, metacarpal I
postorbital bar and jugular bar: absent (0); present fused onto carpus and divergent from rest of manus (0);
(1). reduced to two small, unfused carpals (1).
86. Infratemporal fenestra: posterior margin of pos- 102. Manus, digit 1: metacarpal and one phalanx
torbital straight to gently curving onto supratemporal present (0); entire digit absent (1).
bar (0); distinct oblique angle between posterodorsal 103. Metacarpal III, relative position of proximal end:
margin and supratemporal bar, creating a boxy aligned with those of metacarpals II and IV (0); offset
infratemporal fenestra (1) (new). distally relative to metacarpals II and IV (1).
87. Cervical vertebrae, number: 11 or fewer (0); 12–15 104. Metacarpal, shape: short and robust, width at
(1). midshaft to length ratio of 0.2 or greater (0); slender
and elongate, width at midshaft to length ratio of 0.15 process (0); shaft short, dorsoventral expansion begins
or less (1). at base of process (1).
105. Penultimate phalanges of digits II and III, shape: 112. Pubis, obturator foramen: closed or partially
rectangular, lateral sides subequal in length (0); closed ventrally by tubercle arising from pubic shaft
wedge-shaped, medial side significantly shorter than (0); fully open, tubercle absent (1).
lateral side (1). 113. Ischium, shape of shaft in lateral view: strongly
106. Ilium, size of supracetabular process: small, curved downward (0); nearly straight (1).
projects only as a lateral swelling (0); large, broadly 114. Ischium, shape of distal end: small knob-like foot
overhangs the lateral side of the ilium and usually (0); large and pendant foot (1).
extends at least halfway down the side of the ilium (1). 115. Ilium, shape of dorsal margin: nearly straight (0);
107. Ilium–pubis articulation: large iliac contribution, distinctly depressed over supracetabular process and

pubic peduncle of ilium long, iliac peduncle of pubis dorsally bowed over base of preacetabular process (1).
small (0); pubic process of ilium short with restricted 116. Femur, development of intercondylar extensor
articular surface, prominent dorsally directed iliac groove: moderately deep, groove fully open (0); deep,
peduncle of pubis (1). edges of groove meet or nearly meet cranially to
108. Ilium, postacetabular process shape: tapers pos- enclose an extensor tunnel (1).
teriorly to nearly a point, wide brevis shelf (0); rectan- 117. Tarsus, distal tarsals 2 and 3: present (0); absent
gular, no brevis shelf (1). (1).
109. Ilium, postacetabular process size relative to 118. Metatarsal I, length: short, thin splint (0); absent
total length of ilium: less than 40% (0); more than 40% (1).
(1). 119. Pes, distal phalanges of pedal digits II to IV:
110. Pubis, distal width of prepubic process: dorsoven- axially shortened to disc-like elements with width at
trally expanded to no more than twice the depth of the least three times length (0); greatly shortened, width
proximal shaft (0); expanded to more than twice the at least four times length (1).
depth of the proximal shaft (1). 120. Pes, shape of unguals: taper evenly distally,
111. Pubis, length of prepubic process constriction: claw-like (0); dorsoventrally flattened and broadened,
long, dorsoventral expansion restricted to distal hoof-like (1).
APPENDIX 2
TAXON CHARACTER STATE MATRIX
5 10 15 20 25 30 35 40 45 50
Igaunodon 00000 10000 00000 00000 00000 00000 00000 00000 00000 00000
Eolambia 100?0 01110 000?? ??00? ?1010 00000 0???? ??0?? ?0?0? ??21
Brachylophosaurus 21110 12112 10111 11111 12011 01110 01011 11011 10001 10?21
Corythosaurus 11111 01111 11111 11111 11010 010?1 0?2?2 00100 0?01? 11121
Edmontosaurus spp. 31110 12112 10111 11111 12111 11120 01001 12000 00000 10?21
Gryposaurus notabilis 21110 12111 11101 11111 11111 01110 11010 11012 11001 10021

G. incurvimanus 21110 12111 111?? ?1111 11111 01110 11010 11011 11000 10021
G. latidens 211?0 ?2111 111?? ??111 11111 01110 11010 11011 11000 10021
G. monumentensis 21110 12111 11101 11111 11111 01110 11010 11011 11001 10021
Lophorothon ??10? 01?1? ????? ????? ????? ????? ????? 11012 10??? ?????
Maiasaura 21110 12112 10111 11111 12011 ?1000 00000 11011 10001 10021
Naashoibitosaurus 2???? ?211? ????? ????? ????? ????0 0?0?? 11012 1010? ?0?21
Prosaurolophus spp. 21110 12111 10111 11111 11101 ?1120 01001 12012 10100 10021
Saurolophus spp. 31110 12111 10111 01111 11000 01120 00101 12012 10100 10021
Telmatosaurus 01011 01100 101?? ?0001 1100? 00000 00000 00000 00000 ?0?21
55 60 65 70 75 80 85 90 95 100
Igaunodon 00000 00000 00010 00000 00000 00000 00??0 00000 00000 00000
Eolambia ??000 ????? ???0? 0?0?? 0??00 ??000 0?000 ????0 ????? ??110
Brachylophosaurus 00001 10201 11110 00111 11111 11001 11001 01011 01111 10111
Corythosaurus 12001 11012 10000 11111 11211 ?11?? 11111 11111 11111 10111
Edmontosaurus spp. 00011 10112 10000 00111 11111 11001 11001 01111 01111 10110
Gryposaurus notabilis 01011 10101 11111 00111 11?11 01001 11001 1???? ????? ?????
G. incurvimanus 00011 10101 11111 00011 11?11 01001 11001 11111 01111 00110
G. latidens 000?1 10101 10111 ????? ????? ????? ????? ????? ????? ?????
G. monumentensis 01011 10101 11111 00011 11111 01001 11??1 1?11? 01111 00110
Lophorothon ??001 00??? ????? 0???1 ???1? ?1??1 0???? 0??? ????? ?????
Maiasaura 00111 10201 11110 01111 11211 01??? 11001 11011 01111 00111
Naashoibitosaurus 02001 10101 10112 1?111 1?111 ?1??? 1?001 1???? ????? ??11?
Prosaurolophus spp. 01101 10112 10002 11111 11111 11011 11001 01111 01111 10110
Saurolophus spp. 00101 10112 10003 11111 11211 11??0 11001 01111 01111 00110
Telmatosaurus 00??1 00200 0?010 00111 1?000 0???0 0?000 1?11? 0??11 0101?
105 110 115 120
Igaunodon 00000 00000 00000 00000

Eolambia ????? 0000? ??110 ?????
Brachylophosaurus 11111 11101 01101 11111
Corythosaurus 11111 11111 01111 11111
Edmontosaurus spp. 11111 11111 01101 11111
Gryposaurus notabilis ????? ????? ????? ?????
G. incurvimanus 11111 11111 01101 11111
G. latidens ????? ????? ????? ?????
G. monumentensis ????? 11111 01101 ?????
Lophorothon ????? ????? ??10? ??1?1
Maiasaura 11111 11111 01101 11111
Naashoibitosaurus ????? ????? ????? ?????
Prosaurolophus spp. 11111 11111 01101 11111
Saurolophus spp. 11111 11100 01101 11111
Telmatosaurus ????? ????? ????? 1????

A New Species of Gryposaurus (Dinosauria - Hadrosauridae) From The Late Campanian Kaiparowits Formation, Southern Utah, USA

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A New Species of Gryposaurus (Dinosauria - Hadrosauridae) From The Late Campanian Kaiparowits Formation, Southern Utah, USA

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Zoological Journal of the Linnean Society, 2007, 151, 351–376.

A new species of Gryposaurus (Dinosauria:

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Received March 2007; accepted for publication July 2007

ADDITIONAL KEYWORDS: biogeography – Cretaceous – dinosaur – hadrosaur – hadrosaurinae –

INTRODUCTION Kritosaurus, and described the species K. incurvi-

paludal and riparian environments (Roberts, Deino &

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Figure 3. Lateral view of Gryposaurus monumenten-

Figure 5. Line drawings of the four Gryposaurus species

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Prosaurolophus and Naashoibitosaurus (NMMNH

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contacts the posterior side of the maxilla dorsal

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rosaurine taxa in that it is subtriangular in general

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Frontal: The morphology of the frontal is conserved

feetensis (Horner, 1992). Posterior to the quadrate

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Pro-otic: The nearly pentagonal pro-otic contacts the

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Laterosphenoid: In anterior view, the laterosphenoid

is composed of a large lateral condyle that contacts

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PALATOQUADRATE COMPLEX Palatine: The palatine unites the dorsally positioned

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ventral lobe is excavated in RAM 6797, revealing two

dentary anteriorly, the angular and splenial medially,

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process, anteroposterior shortening of the parietal

Observation of the mandibular foramen in other

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change from a subcircular to an oval mandibular

Overall, the general skull morphology of G. monu-

G. monumentensis is in fact boxier than that of G. no-

Utah taxon. However, this ratio is less than that

depth to length ratio of 0.94 (Table 1).

Gryposaurus monumentensis can be further distin-

robust nature of the skull and lower jaws, a feature

that is perhaps best revealed in the predentary,

tions for generating and absorbing heightened bite

angled premaxillae and shorter skull length relative

vertical bite action and increased the bite force of the

increased attachment area for the lower rhamphoth-

Max. skull depth (MSD) (cm)

eca. Whether or not these characters indicate a

Av. ITF area (ITFA) (cm2)

tougher, more fibrous diet relative to that of closely

of a future functional analysis. Nonetheless, given the

distinction in predentary shape between Gryposaurus

species and other hadrosaurid taxa, members of this

ONTOGENETIC CONSIDERATIONS the subject of a separate study, more closely