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Iguanodon bernissartensis

Reptilia -

Taxonomy
Iguanodon bernissartensis was named by Boulenger (1881). Its type specimen is IRSNB 1534, a skeleton, and it is not a trace fossil. Its type locality is Sainte-Barbe coal mine, Bernissart, which is in a Barremian/Aptian lacustrine - small claystone/bituminous coal in the Sainte-Barbe Clays Formation of Belgium.

It was synonymized subjectively with Iguanodon mantelli by Hooley (1912) and Kuhn (1964).

Synonyms
Synonymy list
YearName and author
1881Iguanodon bernissartensis Boulenger
1882Iguanodon bernissartensis Dollo
1882Iguanodon seelyi Dollo
1882Iguanodon bernissartensis Dollo p. 205
1882Iguanodon seelyi Hulke p. 144 figs. Pl. 4
1882Iguanodon seelyi Seeley p. 371
1883Iguanodon bernissartensis Dollo figs. Pl. III
1883Iguanodon bernissartensis Meunier p. 338
1883Iguanodon bernissartensis Moseley p. 440
1883Iguanodon seeleyi Moseley p. 440
1883Iguanodon seelyi Seeley p. 60
1884Iguanodon bernissartensis Dames p. 186
1884Iguanodon bernissartensis Dollo p. 2
1884Iguanodon bernissartensis Hulke p. 48
1885Iguanodon bernissartensis Dollo p. 26
1885Iguanodon bernissartensis Woodward pp. 13-14
1886Iguanodon bernissartensis Cope p. 153
1886Iguanodon bernissartensis Günther p. 442
1887Iguanodon bernissartensis Dollo p. 305
1887Iguanodon bernissartensis Seeley p. 80
1887Iguanodon seelyi Seeley p. 82
1888Iguanodon bernissartensis Lydekker p. 46
1888Iguanodon bernissartensis Mansel-Pleydell p. 34
1889Iguanodon bernissartensis Lydekker p. 355
1890Iguanodon bernissartensis Gaudry p. 212
1890Iguanodon bernissartensis Lydekker p. 36
1890Iguanodon bernissartensis Woodward and Sherborn p. 242
1890Iguanodon bernissartensis Zittel p. 696
1892Iguanodon bernissartensis Dupont p. 89
1893Iguanodon bernissartensis Lydekker p. 304
1894Iguanodon bernissartensis Struckmann p. 829
1895Iguanodon bernissartensis Kehrer p. 355
1895Iguanodon bernissartensis Marsh p. 687
1895Iguanodon bernissartensis Woodward pp. 289-291
1896Iguanodon bernissartensis Anonymous p. 94
1896Iguanodon bernissartensis Gaudry p. 94
1896Iguanodon bernissartensis Harmer p. 202
1896Iguanodon bernissartensis Mansel-Pleydell p. 119
1896Iguanodon bernissartensis Marsh p. 230 figs. Pl. 85
1897Iguanodon bernissartensis Anonymous p. 4
1898Iguanodon bernissartensis Woodward p. 206
1898Iguanodon seelyi Woodward p. 208
1899Iguanodon bernissartensis Savalle and Lennier p. 61
1900Iguanodon bernissartensis Nopcsa p. 562
1902Iguanodon bernissartensis De Pauw p. 85
1902Iguanodon bernissartensis Mansel-Pleydell p. lxxxiii
1902Iguanodon bernissartensis Nopcsa pp. 151-152
1902Iguanodon bernissartensis du Pauw p. 20
1907Iguanodon bernissartensis Seitz p. 325
1908Iguanodon bernissartensis Joubert p. 269
1911Iguanodon bernissartensis Drevermann p. 97–98
1911Iguanodon bernissartensis von Zittel p. 290 figs. 436-437
1913Iguanodon bernissartensis Tornier p. 370 fig. 51
1915Iguanodon bernissartensis Nopcsa p. 385
1917Iguanodon bernissartensis Hooley p. 148
1925Iguanodon bernissartensis Gilmore p. 387
1925Iguanodon bernissartensis Hooley p. 3
1930Iguanodon bernissartensis Osborn p. 531–532
1933Iguanodon bernissartensis Janensch p. 232
1933Iguanodon bernissartensis Swinton p. 66
1934Iguanodon bernissartensis Gross p. 759
1934Iguanodon bernissartensis Swinton p. 41
1936Iguanodon bernissartensis Swinton pp. 214-215
1939Iguanodon bernissartensis Jackson p. 67
1944Iguanodon bernissariensis Young p. 197
1960Iguanodon bernissartensis Lapparent p. 183
1962Iguanodon bernissartensis Lapparent p. 20
1964Iguanodon bernissartensis Tatarinov p. 559 fig. 605
1967Iguanodon bernissartensis Lapparent p. 10
1968Iguanodon bernissartensis Lapparent and Stchepinsky p. 1372
1969Iguanodon bernissartensis Quinet p. 30
1970Iguanodon bernissartensis Swinton p. 209
1974Iguanodon bernissartensis Langston p. 89
1976Iguanodon bernissartensis Antunes p. 21
1978Iguanodon bernissartensis Lehmann p. 109
1979Iguanodon bernissartensis Jurcsak and Popa p. 52
1980Iguanodon bernissartensis Buffetaut et al. p. 201
1980Iguanodon bernissartensis Norman p. 13
1981Iguanodon bernissartensis Bartholomai and Molnar p. 343
1981Iguanodon seelyi Bartholomai and Molnar p. 343
1981Iguanodon bernissartensis Santafé et al. p. 48
1982Iguanodon bernissartensis Santafé et al. p. 99
1983Iguanodon bernissartensis Blows p. 311
1983Iguanodon bernissartensis Weishampel and Weishampel p. 44
1984Iguanodon bernissartensis Buscalioni and Sanz p. 14
1984Iguanodon bernissartensis Sanz p. 503
1984Iguanodon bernissartensis Sanz et al. p. 209
1986Iguanodon bernissartensis Sanz p. 83
1987Iguanodon bernissartensis Buscalioni and Sanz p. 66
1987Iguanodon bernissartensis Casanovas et al. p. 102
1987Iguanodon bernissartensis Norman p. 165
1989Iguanodon bernissartensis Frances and Sanz p. 141
1989Iguanodon bernissartensis Weishampel and Bjork p. 64
1990Iguanodon bernissartensis Norman and Weishampel p. 530
1991Iguanodon bernissartensis Weishampel et al. p. 203
1992Iguanodon bernissartensis Cheevely and Chapman p. 347
1992Iguanodon bernissartensis Martin and Buffetaut p. 68
1994Iguanodon bernissartensis Lockley p. 350
1996Iguanodon bernissartensis Molnar p. 640 fig. 2
1997Iguanodon bernissartensis Buffetaut p. 214
1997Iguanodon bernissartensis Currie and Padian p. 369
1997Iguanodon bernissartensis Forster p. 360
1997Iguanodon bernissartensis Norman p. 784
1998Iguanodon bernissartensis Blows p. 31
1998Iguanodon bernissartensis Godefroit et al. p. 47
1998Iguanodon bernissartensis Ruiz-Omeñaca et al. p. 275
1999Iguanodon bernissartensis Cuenca-Bescós et al. p. 237
1999Iguanodon bernissartensis Pereda Suberbiola et al. p. 271
2000Iguanodon bernissartensis Norman and Sues p. 463
2001Iguanodon bernissartensis Wang and Xu p. 1671
2003Iguanodon bernissartensis Godefroit and Knoll p. 5
2003Iguanodon bernissartensis Posmosanu p. 432
2003Iguanodon bernissartensis Ruiz-Omeñaca and Canudo p. 275
2004Iguanodon bernissartensis Norman p. 416
2005Iguanodon bernissartensis You et al. p. 786
2006Iguanodon bernissartensis Pereda Suberbiola et al. p. 227
2006Iguanodon bernissartensis Prieto-Márques et al. p. 936
2007Iguanodon bernissartensis Brill and Carpenter p. 60
2007Iguanodon bernissartensis Contreras et al. p. 53
2007Iguanodon bernissartensis Paul p. 74
2007Iguanodon bernissartensis Sánchez-Hernández et al. p. 206
2008Iguanodon bernissartensis Naish and Martill p. 617
2008Iguanodon bernissartensis Paul p. 199
2008Iguanodon bernissartensis Ruiz-Omeñaca et al. p. 65
2009Iguanodon bernissartensis Kim et al. p. 1394
2010Dollodon seelyi Carpenter and Ishida p. 148
2010Iguanodon bernissartensis Carpenter and Ishida p. 148
2010Iguanodon bernissartensis McDonald et al. p. 2
2010Iguanodon bernissartensis McDonald et al. p. 33 fig. 39
2010Iguanodon bernissartensis McDonald et al. p. 808 fig. 10
2010Iguanodon bernissartensis Norman p. 64
2010Iguanodon bernissartensis Pereda Suberbiola et al. p. 64
2010Iguanodon bernissartensis Prieto-Márquez p. 3
2010Iguanodon bernissartensis Prieto-Márquez and Wagner p. 1243 fig. 3
2011Iguanodon bernissartensis Gasulla et al. p. 160
2011Iguanodon bernissartensis Gates et al. p. 808 fig. 12
2011Iguanodon bernissartensis Pereda-Suberbiola et al. p. 556
2011Iguanodon bernissartensis Prieto-Márquez p. 63
2011Iguanodon bernissartensis Wang et al. p. 137
2012Iguanodon bernissartensis Han et al. p. 1370
2012Iguanodon bernissartensis Prieto-Marquez et al. p. 31
2012Iguanodon bernissartensis Ramírez-Velasco et al. p. 390
2013Iguanodon bernissartensis Llandres Serrano et al. p. 573
2013Iguanodon bernissartensis Norman p. 395–397
2013Iguanodon bernissartensis Pereda Suberbiola and Ruiz-Omeñaca pp. 13, 15
2013Iguanodon bernissartensis Zhang et al. p. 1346
2013Iguanodon bernissartensis Zheng et al.
2014Iguanodon bernissartensis Gasulla et al. p. 19
2014Iguanodon bernissartensis McDonald et al. p. 46
2014Iguanodon bernissartensis Norman p. 22
2014Iguanodon bernissartensis Sanz et al. p. 76
2015Iguanodon bernissartensis Verdú et al. p. 251
2018Iguanodon bernissartensis Madzia et al. p. 968

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RankNameAuthor
kingdomAnimalia()
Triploblastica
Nephrozoa
Deuterostomia
phylumChordataHaeckel 1847
OlfactoresJefferies 1991
subphylumVertebrata
Gnathostomata()
Osteichthyes()
Sarcopterygii
subclassDipnotetrapodomorpha(Nelson 2006)
subclassTetrapodomorpha()
Tetrapoda()
Reptiliomorpha
Anthracosauria
Batrachosauria()
Cotylosauria()
Amniota
Sauropsida
classReptilia
subclassEureptilia()
Romeriida
Diapsida()
RankNameAuthor
Eosuchia()
Neodiapsida
SauriaGauthier 1984
Archosauromorpha(Huene 1946)
Crocopoda
ArchosauriformesGauthier 1986
Eucrocopoda
Archosauria()
informalAvemetatarsalia
Ornithodira
Dinosauromorpha
Dinosauriformes
Dinosauria()
Ornithischia()
Neornithischia()
Ornithopoda()
Iguanodontia()
Dryomorpha
Ankylopollexia
Styracosterna
Hadrosauriformes
genusIguanodon
speciesbernissartensis

If no rank is listed, the taxon is considered an unranked clade in modern classifications. Ranks may be repeated or presented in the wrong order because authors working on different parts of the classification may disagree about how to rank taxa.

Diagnosis
ReferenceDiagnosis
L. Dollo 1882Skull twice as long as wide, measuring around 65 centimeters. Nares are projected over only one-third of the lower jaw. Greater diameter of the orbit vertical. Temporal fossa largely open. Minimum width of the scapula contained nearly six times in its maximum length. Coracoid furnished with an indentation communicating laterally with the exterior. Sternal plates wide and strongly interiorly indented. Forelimbs representing nearly two-thirds of the hindlimbs. Six vertebrae in the sacrum. Preacetabular projection of the ilium equal to about one-third of the total length of this bone. Pubis long, massive, and straight. Third trochanter situated near the lower third of the femur. Total length 8 to 10 meters.
D. B. Norman 1980Maximum length 11 metres. Double supraorbitals (palpebrals). Number of vertical tooth rows in maxilla and dentary variable, with a maximum of 29 in the maxilla an 25 in the dentary. Sacrum composed of eigth fused vertebrae. Scapula, inexpanded distally, proximal region of shaft considerably thickened. Intersternal ossification present. Forelimb long (approx. 70% of length of hindlimb) and stroutly constructed. Carpals co-ossified. First metacarpal fused to carpals. Phalangeal count of manus: 2, 3, 3, 4, 4; first phalanx of digit I consists of a thin, warped plate of bone, which lies in a shallow recess in the proximal surface of the ungual phalanx. The latter has the form of a long, curved, conical spine which can freely articulate against the fused carpometacarpus. Ungual phalanges of digits II and III are broad, rather hoof-like and twisted longitudinally. Hind end of ilium pointed in lateral aspect, with a broad brevis shelf. Anterior ramus of the pubis relatively narrow, proximally transversely flattened, but with a moderately expanded distal end. Three distal tarsals present. Metatarsal I reduced and transversely flattened, matatarsal V absent.
G. S. Paul 2007Adults large at 11 m, mass about 3+ tonnes. Overall massively constructed. Accessory palpebral present. Quadrate shaft straight. Maximum tooth positions 29 in maxila, 25 in dentary. Neural spines of dorsals, sacrals, and caudals short. Posterior dorsal centra anteroposteriorly compressed. Eight fused sacrals. Scapula consistently broad along length of blade. Acromion process placed proximally and directed anteriorly. Coracoid large relative to scapula. Forelimb 75% of hindlimb. Deltopeectoral crest of humerus modest sized. Manual spike and other unguals large. Prepubis shallow. Femoral fourth trochanter ventrally spiked.
G. S. Paul 2008Overall massively constructed. Premaxillary tip to anterior orbital rim/latter to paraoccipital process tip length ratio ~1.1; dentary pre-coronoid process length/minimum depth ratio under 4. Maxillary process of premaxilla shallow. Dorsal apex of maxilla set posteriorly. Antorbital fossa and fenestra *moderate in size. Lacrimal short, does not contact nasal. Accessory palpebral *present. Posterior border of occiput deeply indented. Lateral temporal fenestra large. Posterior portion of jugal short. Quadratojugal tall. Quadrate tall, transversely broad, shaft straight, lateral foramen set low, dorso-posterior buttress *very large. Diastema absent. Maximum tooth positions *29 in maxilla, *25 in dentary. Dorso-sacral/hindlimb length ratio w1. Posterior dorsal centra antero-posteriorly compressed. Neural spines of dorsals, sacrals and caudals short. *8 fused sacrals. Scapula blade *consistently broad, *base very broad, *acromion process placed dorsally and directed anteriorly. Coracoid large. Forelimb *~70% length of hindlimb. Deltopectoral crest of humerus *proximally located and modest in size. Olecranon process large. Manus *massive, phalanx 1 of digit I *present, pollex spike and other unguals large. Anterior process of ilium *much shorter than main body, main body shallow, posterior acetabular body *long. Prepubic process of pubis shallow. Femoral shaft straight. Metatarsal I present, II long.
F. J. Verdú et al. 2015Potential autapomorphies of I. bernissartensis according to Norman (2012): (I) two palpebral bones; (II) 29 maxillary and 25 dentary teeth; (III) posterior dorsals remain tall-sided, but broaden to produce an almost circular shape (indented dorsally by the neural canal), giving rise to a broadly heart-shaped articular face when viewed axially; (IV) scapular blade tends to be comparatively slightly expanded distally and modestly recurved in lateral view (...), with the proximal portion of the shaft expanded and massive adjacent to the scapulocoracoid suture and humeral glenoid; (V) an “intersternal ossification” is commonly preserved in the midline of the upper chest in the space between the coracoids and sternal bones; (VI) forelimb at least 70% the length of the hind limb, and robustly constructed; (VII) carpals heavily co-ossified and supported by ossified ligaments; (VIII) phalangeal count of the manus: 2:3:3:2:4; (IX) the first phalanx of digit 1 of the manus is preserved as a flattened and warped plate that lies in a recess at the base of the pollex ungual; (X) united phalanx 1–pollex can articulate freely (and transversely) against the roller-like articular surface of metacarpal 1; (XI) three distal tarsals are present; (XII) metatarsal I reduced to a transversely flattened splint that lies obliquely against the shaft of metatarsal II.
An additional potential autapomorphy of I. bernissartensis might be an ischium with a distal end small compared to its proximal end (ratio 0.27 in IRSNB 1534 [holotype] in Fig. 67 of Norman (1980)).