22
84 EXPLANATION OF PLATE I. PLATE I. - lguanodon bernissartensis. IRSNB. 1561. Partly disarticulated skull with lower jaw. Redrawn from photographs.
84
EXPLANATION OF PLATE I.
PLATE I. - lguanodon bernissartensis. IRSNB. 1561. Partly disarticulated skull with lower jaw. Redrawn from photographs.
Inst. Roy. Sc. Nat. Belg. - Mem. N° Kon. Belg. Inst. Natuurw. - Verh. Nr
D.B. NORMAN. - On the ornithiscbian dinosaur Jguanodon bernissartensis from the Lower Cretaceous of Bemissart (Belgium)
PI.I
86
EXPLANATION OF PLATE II.
PLATE II. - lguanodon bernissartensis. IRSNB. 1561. Partly disarticulated skull with lower jaw. Redrawn from photographs.
Inst. Roy. Sc. Nat. Belg. - Mem. N° Kon. Belg. Inst. Natuurw. - Verh. N'
D.B. NORMAN. - On the ornithischian dinosaur
E (.)
0 .....
/guanodon bernissartensis from the Lower Cretaceous of Bernissart (Belgium)
Pl. II
87
88
EXPLANATION OF PLATE III.
PLATE III. - Jguanodon bernissartensis. IRSNB. 1536. Partly disarticulated skull with lower jaw. Redrawn from photographs.
Inst. Roy. Sc. Nat. Belg. - Mem. N° Kon. Belg. Inst. Natuurw. - Verb. N '
D .B. NORMAN.·_ On the ornithischian dinosaur lguanodon bernissartensis from the Lower Cretaceous of Bernissart (Belgium)
Pl. III
89
90
EXPLANATION OF PLATE IV.
PLATE IV. - Jguanodon bernissartensis. IRSNB. 1536. Partly disarticulated skull with lower jaw. Redrawn from photographs.
Inst. Roy. Sc. Nat. Belg. - Mem. N° Kon. Belg. Inst. Natuurw. - Verh. N'
D.B. NORMAN. - On the ornithischian dinosaur /guanodon bernissartensis from the Lower Cretaceous of Bernissart (Belgium)
Pl. N
91
92
EXPLANATION OF PLATE V.
PLATE V. - Part of a trackway of footprints which was until recently displayed in the cc Dinosaur gallery » of the British Museum (nat. Hist.). The trackway comprises two nearly parallel sets of footprints, one of which exhibits characteristic << megalosaur >> prints, the other probably belongs to lguanodon.
Inst. Roy. Sc. Nat. Belg. - Mem. N° Kon. Belg. Inst. Natuurw. - Verh. N'
A
~l en c 0 en en Q) ... Cl. E
0 0 ..... Q) ... 0
LL
D.B. NORMAN. - On the ornithischian dinosaur lf!,uanodon bernissartensis from the Lower Cretaceous of Bernissart (Belgium)
Pl. V
93
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95
DAVID BRUCE NORMAN - ON THE ORNITHISCHIAN DINOSAUR IGUANODON
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96
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BERNISSARTENSIS FROM THE LOWER CRETACEOUS OF BERNISSART (BELGIUM)
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188 1, Sur I' arc pelvien clzez les dinosauriens de Bernissart. (Bull. Acad. R. Beige. Ch. Sci., (3) l : 600-608.)
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97
G - INDEX OF ABREVIATIONS
A articular J ju gal a.zyg prezygapophysis ac = acetabulum k keel ad.f adductor fossa An angular L lachrymal an.pu prepubic ramus 1.tr lesser trochanter ao antorbital foramen li ligament scars Ap alveolar parapet lip lateral lip on worn dentary teeth art articulating surface l.M lateral malleolus (tibia) as astragalus
M maxilla as.p ascending process (astragalus) me metacarpal asc ascending process (supraoccipital) Me Meckelian canal at atlas
ax axis mt metatarsal
N nasal b boss (scapula) n.ca neural canal bi blunt ended (ribs) nc.s neuro-central suture Bo basioccipital ns neural spine Bpt basipterygoid process
Bs basisphenoid ol olecranon Op opisthotic c carpal op.s opisthotic suture
c capitulum Os orbitosphenoid
c. for coracoid foramen os.ligs ossified ligaments ea calcaneurn
ot ossified tendons Ca.r caudal rib
obt obturator process Carp carpal bone eh chevron (haemal arch) p parietal en cnemial crest p primary ridge Co coronoid p .at proatlas cor.s = coracoid suture p .pu post-pubic ramus
p.zyg postzygapophysis D dentary Pa prearticular d = diapophysis pa parapophysis D (Roman numeral) = digit number Pd predentary d.cr de! to-pectoral crest Pf prefrontal
Ph phalanx E exoccipi tal Pm premaxilla e.s exoccipital suture Po postorbital Ee ectopterygoid
Pr proi.itic Ec.p ectopterygoid ramus pr process
Ps parasphenoid F frontal Pt pterygoid F.te flexor tendon scar pt.f post-temporal foramen fo fenestra ovalis Pt/ Q pterygoid and quadrate flanges for foramen (foramina) pt.s pterygoid suture pu pubis g. tr greater trochanter pu.p pubic peduncle gl glenoid paf parietal fissure gr grooves Pal palatine
h head (squamosal) Pap paroccipital process h.s haemal sulcus pit area of pituitary fossa he heel pos postorbital suture hr horizontal ridge pro proximal phalanx
pyr pyrites ig intercondylar groove
Q quad rate il = ilium Q-j quadrato-jugal is ischium Qr quadrate ramus is.Ii ligament scars on pubis (ischial)
is.p ischial tuber r.c replacement crown int intermedium ra radius
99
DA YID BRUCE NORMAN
re resorption facet ri rib spine ra.f radial facet ro.c root-less crown ru.par rugose parapophysis rad radiale rug rugosities rad.c radial condyle
s splenial s secondary ridge s.s scapular suture S.sh splenial shelf Sa surangular SC = muscle scar sh = shaft (of rib) So supraorbital - (palpebral) Sq squamosal Su supraoccipital sa.r sacral rib sp.b spongy bone sq.b = squamosal « boss D
100
ON THE ORNITHISCHIAN DINOSAUR IGUANODON
sq.p squamosal process St.hi " standard hindlimb » sep separation of pedicels of haemal arch
tertiary ridge t (Roman numeral = distal tarsal te tooth crowns ti = tibia tp transverse process tu tuberculum
ul ulna ul.c ulnar condyle ul.f ulnar facet uln ulna re Ung ungual phalanx
V vomer v.for vcd
w 4th.tr
vascular foramina = canal of vena capitis dorsalis (post-temporal
foramen) wear facet fourth trochanter
TEXT - FIGURE LEGENDS
FIG. 1. - An outline map of Belgium to illustrate the locations of Bernissart
FIG. 2. - lg11a11odo11 bemissarte11sis. Skull reconstruction, based mainly on: IRSNB. 1561, 1536, 1535. For abreviations see alphabetical index, section G ...
FIG. 3. - l g11anodo11 bernissartensis. Skull reconstruction in dorsal view ...
FIG. 4. - l g11a11odo11 bernissartensis. Skull reconstruction. A, anterior view; B, posterior (occipital) view. Broken lines indicate imperfectly known regions ...
FIG. 5. - l g11a11odo11 bemissartensis. Skull reconstruction, in ventral view, lower jaws removed. Broken Jines indicate imperfectly known regions .. .
FIG. 6. lg11a11odo11 bemissartensis. The occipital region of A, IRS B. 1535 (en gisement; B, IRS B. 1561 (Monte) ...
FIG. 7. - The supraoccipital of an unidentified hadrosaur. Redrawn from Langston (1960) Plate 34, Figs. B, E ....
FIG. 8. - l g11a11odo11 hemissartensis. IRS B. 1536. Postero-medial view of the occipital region of the skull. Cross-hatching indicates broken bone surface; even stippling indicates longitudinal section ...
FIG. 9. - l g11a11odo11 bemissartensis. Reconstruction of the skull which has been partly dissected to reveal the content of the naso/buccal cavities and the structures of the braincase. Detailed structure of braincase from /. mantelli (BM H. R. 2501.). Even stippling represents • cut• bone surface
FIG. 10. - l g11a11odo11 bemissartensis. (IRS B. 1536). Orbital region of the skull. Broken line indicates approximate area of suture between Frontal and Postorbital .
FIG. 11. - l g11a11odo11 bemissartensis. (1RSNB. 1536). Right Supraorbital bones. A : dorsal view; B : medial view; C : ventral view; D : lateral view ...
FIG. 12. - l g11a11odon bemissartensis. (1RSNB. 1535). Left Quadrate and attached portions of the Jugal, Quadrato-jugal and pterygoid, as preserved ...
FIG. 13. - lg11a11odo11 bemissartensis. (IRS B. 1536). Posterior portion of the left half of the skull. Quadrato-jugal i11 si111 ...
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18
18
19
20
21
22
23
24
FIG. 14. - Iguanodon bernissartensis. (BMNH. R . 8306). Ventral view of the palatal region of a partial skull. Even stippling indicates matrix, cross-hatching indicates broken bone surface .. .
FIG. 15. - lguanodon bernissartensis. (IRSNB. 1561). Predentary. Dorsal view of left half, as preserved, articulated with the D entary
FIG. 16. - Ig11a11odon bernissartensis. Posterior view of the hind end of the lower jaw. A : IRSNB. 1561; B: IRSNB. 1536 . ..
FIG. 17. - /guanodon bemissartensis. (IRSNB. 1561). Hyoid bone. A : lateral view; B : medial view
FIG. 18. - Iguanodon bernissartensis. (IRSNB. 1561). Selected dentary teeth. A, B: anterior and medial views of an anterior dentary tooth as indicated in fig. 19. C : occlusal view of teeth indicated in fig. 19. D , E : la teral and occlusal views of teeth as indicated in fig. 19
FIG. 19. - Iguanodon bemissartensis. (lRSNB. 1561). Dentition of the left lower jaw, as preserved. Replacement • wave • indicated by even tone ...
FIG. 20. - l guanodon sp. (BM H . R. 40100). Selected maxillary teeth, demonstrating replacement process in si111 ...
FIG. 21. - l guanodon bemissarlensis. (IRSNB. 1536). Maxillary dentition, anterior portion, as preserved. Replacement • wave • indicated by even stippling
FIG. 22. - lguanodon bemissartensis. The artic-ulated cervical vertebrae. Based on IRSNB. 1536, 1561 ...
FIG. 23. - lguanodon bemissartensis. (IRSNB.
FIG. 24.
FIG. 25.
1561). Atlas vertebrae as preserved. a : lateral view; b : dorsal view; c: ventral view; d : anterior view ...
- /g11anodon bemissartensis. (IRSNB. 1561). Axis vertebra and odontoid, as preserved. a : lateral view; b : posterior view; c: ventral view
- lguanodon bernissartensis. (IRSNB. 1561). 3rd Cervical vertebra, as pre-served. a: lateral view; b: posterior view; c : dorsal view ...
FIG. 26. - l g11a11odo11 bemissartensis. (IRS B. 1561). 6th Cervical vertebra, as preserved. a : lateral view; b : ventral view; c : dorsal view ...
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DA YID BRUCE NORMAN - ON THE ORNITHISCHIAN DINOSAUR IGUANODON
FIG. 27. - Ig11anodon bernissartensis. (IRSNB. 1561). Atlas rib (right) medial view ...
FIG. 28. - Jguanodon bernissartensis. (IRSND. J 561). Axis rib, (left). a : lateral ivew; b : medial view ...
FIG. 29. - Ig11a11odon bernissartensis. (IRSNB. 1561). 3rd Cervical rib (right). a : medial view; b : lateral view
FIG. 30. - Iguanodon bernissartensis. (IRSNB. I 536). 4th Cervical rib (right). a : medial view; b : lateral view
FIG. 31. - Iguanodon bernissartensis. (IRSNB. 1536). 5th Cervical rib (right). a: medial view; b : lateral view
FIG. 32. - lguanodon bernissartensis. (IRSNB. 1536). 8th Cervical rib (right) lateral view
FIG. 33. - lguanodon bernissartensis. (IRSNB. 1536). !Oth Cervical rib (left) lateral view
FIG. 34. - Iguanodon bernissartensis. Reconstruction of the anterior dorsal vertebral column. Based on « individu S » of the conservatoire, l.A.Sc.N .B. Lateral view
FIG. 35. - lguanodon bernissarte11sis. Reconstruction of the anterior dorsal vertebral column as in fig. 34. A : dorsal view; B : ventral view . ..
FIG. 36. - l guanodon bernissartensis. (« lndividu S •) a, b : 2nd Dorsal vertebra in lateral and anterior views; c, d : 5th Dorsal vertebra in lateral and anterior views ...
FIG. 37. - l guanodon bernissartensis. Reconstruction of the posterior dorsal vertebral column, based on « lndividu S », vertebrae 14 ·and 16 have been restored. Lateral view
FIG. 38. - l guanodon bernissartensis. Reconstruction of the posterior dorsal vertebral column as in fig. 37. A : dorsal view; B : ventral view ...
FIG. 39. - Iguanodon bernissartensis. (« Individu S •). a: posterior view of 6th Dorsal vertebra; b, c, d: 15th Dorsal vertebra in lateral, anterior and posterior views respectively
FIG. 40. - Iguanodon bemissartensis. (Holotype. IRSNB. 1534). The articulated dorsal vertebral column, as preserved. A : lateral view; B : ventral view
FIG. 41. - lguanodon bernissartensis. (IRSNB. 1536). I ste Dorsal rib (left) in anterior view
FIG. 42. - lguanodon bernissartensis. A typical anterior dorsal rib ,anterior view
FIG. 43. - lguanodon bernissartensis. (Holotype.
102
IRSNB. 1534). a, b: 14th Dorsal rib in dorsal and ventral view respectively. c, d: 16th Dorsal rib in dorsal and ventral view respectively
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FIG. 44. - Ig11anodo11 bemissartensis. (IRSNB. 1729). The Sacrum. Lateral view as preserved. Sacral rib yoke emphasised by even stippling, broken bone crosshatched ..
FIG. 45. - Jg11a11odo11 bemissartensis. The sacrum. A: IRSNB. 1722, as preserved, in ventral view; B : JRSNB. 1535, as preserved, in ventral view ...
FIG. 46. - Jguanodon bemissartensis. (IRSNB. 1729). The articulated tail vertebrae, as preserved « en gisement » ...
FIG. 47. - Ig11a11odo11 bernissartensis. (Holotype. JRSNB. 1534). Articulated series of anterior caudal vertebrae (1-9). Even stippling indicates « carton pierre • restoration
FIG. 48. - lguanodon bemissartensis. (IRSNB. 1725). Posterior caudal vertebrae, as preserved •en gisement ». Cross-hatching represents broken surfaces
FIG. 49. - lg11a11odon bemissartensis. (lRSNB. 1715). Posterior caudal vertebrae, as preserved «Monte • ...
FIG. 50. - lgua11odon bernissarrensis. (I RSNB. 1561). Chevron bone (haemal arch) as preserved. a : left lateral view; b : anterior views; c : posterior view ...
FIG. 51. - Jg11a11odo11 bemissartensis. (IRS B. 1561). Ossified tendons as preserved on the neural spines of the posterior dorsal vertebrae. Right lateral view ...
FIG. 52. - lg11a11odo11 bemissartensis. Scapula (left). A: lateral view (Holotype. lRSNB. 1534) as preserved; B : posterior view (restored); C: medial view (restored). Even stippling represents areas restored with « carton pierre •
FIG. 53. - lguanodon bemissartensis. (Holotype. IRSNB. 1534). Coracoid (left). A : lateral view; B : posterior view; C : medial view; D : anterior view
FIG. 54. - Iguanodon bernissartensis. (Holotype. JRSNB. 1534). Sternal bone (left). A : internal view; B : exteral view. Even stippling represents areas restored with « carton pierre • .
FIG. 55. - lg11anodo11 bemissartensis. (IRSNB. 1535). Sternal region of an articulated skeleton « en gisement • ...
FIG. 56. - Jguanodon bemissartensis. (JRSNB. 1536). Sternal region of a mounted skeleton, as preserved .
FIG. 57. - lguanodon bernissartensis. . (Holotype. JRSNB. 1534). Humerus (right). A: dorsal surface; B : ventral surface. Even stippling - « carton pierre »; cross-hatching - broken bone surface
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BERNISSARTENSIS FROM THE LOWER CRETACEOUS OF BERNISSART (BELGIUM)
FIG. 58. - l guanodon bemissartensis. (Holotype. IRSNB. 1534). Radius and ulna (right). A : medial view; B : lateral view. Even stippling - "carton pierre •
FIG. 59. - l g11anodo11 bemissartensis. (IRSNB. 1558). Articulated carpal bones. a : distal surface; b : dorsal surface; c : proximal surface; d : ventral surface ...
FIG. 60. - Iguanodon bernissartensis. (JRSNB. Articulated digits. a : dorsal ; b : ventral. Even stippling - cc carton pierre •; crosshatching - broken bone surface .. .
FIG. 61. - l g11a11odo11 bernissartensis. (Holotype. JRSNB. 1534). Articulated carpalia and digit 1 in : a: dorsal; b: ventral view. Articulated digits II - IV in : c: dorsal; d : ventral view. e-h : Well preserved first phalanx of digit 1 found in conservatoire collection « Individu H •. e: distal view; f : medial view; g : proximal view; h : lateral view ...
FIG. 62. - /g11anodon bemissa/'/e11sis. The third phalanx of Digit IV ...
FIG. 63. - Ig11anodon bernissartensis. (Holotype. IRSNB. 1534). Ilium (left). Lateral view
FIG. 64. - l guanodon bernissartensis. Pelvic girdle reconstruction, based mainly on lRSNB. 1534
FIG. 65. - l g11anodo11 bernissartensis. (Holotype. lRSNB. 1534). Pubis (left) lateral view.
FIG. 66. - Iguanodon bemissarre11sis. (IRSNB. 156 1). Medial view of the proximal end of the post-pubic ramus of the left pubis
FIG. 67. - Jguanodon bemissar1e11sis. (Holotype. IRSNB. 1534). Ischium (right). A : lateral view; B : medial view ...
FIG. 68. - l g11a11odon bemissartensis. (Holotype. IRSNB. 1534). Femur (right). a : anterior (dorsal) view; b : medial view; c : posterior (ventral) view; d : lateral view .
FIG. 69. - Iguanodon bernissarrensis. (Holotype. JRSNB. 1534). Articulated cruro-tarsus. a : anterior (dorsal) vieww; b : medial view; c : posterior (ventral) view; d : lateral view ...
FIG. 70. - Ig11anodo11 bemissarlensis. (Holotype. lRSNB. 1534). Articulated distal tarsals and metatarsals. A : anterior (dorsal) view; B : posterior (ventral view); C:: proximal view; D : medial view ...
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FIG. 71. - Iguanodon bernissartensis. (Holotype. IRSNB. 1534). Pes (right). As preserved (from photograph) .
FIG. 72. - Iguanodon bernissartensis. (Holotype. lRSNB. 1534) Digit llI of pas. A: lateral; B : ventral; C: dorsal .
FIG. 73. - The prehensile tongue of lguanodon. Redrawn from HEILMANN (1928, fig. 3) ...
FIG. 74. - A reconstruction of l guanodon made by Gideon MANTELL (circa 1835) ...
FIG. 75. - A reconstruction of l guanodon bernissartensis. Redrawn from D OLLO (l 882a)
FIG. 76. - l guanodon bernissartensis. (IRSNB. 1561). and / . mantelli (IRSNB. 1551) as preserved in situ. Redrawn from original archive material. Originals by G . LAVALETTE
FIG. 77. - Iguanodon man1elli. The articulated left man us, based on BMNH. R. 5764 ...
FIG. 78. - l guanodon bernissartensis. (IRSNB. 1558). The articulated left manus in an hyperextended (weight-bearing) position. Redrawn from a photograph ...
FIG. 79. - l guanodon mantelli. (Based on BMNH.
FIG. 80.
R. 5764). The left manus in a weightbearing position ...
lg11a11odo11 bemissarlensis. (IRSNB. 1558). The left manus in a flexed (grasping) position .. .
FIG. 81. - Comparison of the relative size and proportions of the forelimbs of selected ornithopods ..
FIG. 82. - Graphical representation of limb proportions in a variety of dinosaurs . ..
FIG. 83. - I. mantelli. An attempt at a complete skeletal restoration in a normal walking pose
FIG. 84. - I. bemissartensis. An attempt at a corn-plete skeletal restoration In normal walking pose
FIG. 85. - l guanodon bemissarte11sis. (IRSNB. 1730). A: the humerus; B : the femur; as preserved
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ABSTRACT .. .
ACKNOWLEOGE~tr TS
A. - INTRODUCTION
B. - HISTORICAL REVIEW
Discovery and Excavation ...
Preparation and Exhibition .
Literature review ..
C. - OSTEOLOGY
Specimens used for description ...
The skull and lower jaw
General description .
Neurocranium
Maxillary segment ...
Mandible ..
Accessory Skull Elements
Dentition ...
The vertebral .::olumn
Cervical vertebrae ..
Cervical ribs ..
Dorsal vertebrae
Dorsal ribs
Sacrum
Caudal vertebrae
Haemal arches (chevrons)
Ossified tendons .. .
H-CONTENTS
P ages
3
5
7
9
9
10
10
13
13
13
15
19
25
27
28
30
33
35
39
40 42
44
44
The pectoral girdle and forelimb
The pelvic girdle and hindlimb ...
D. - ANATOMICAL OBSERVATIONS ..
Cranial
The horny beak ...
The tongue
Supraorbitals (palpebrals)
The occiput
Postcranial
Natural posture of I . bernissartensis and I . mantelli ...
Forelimb function in / . bemissartensis and I. mantelli ...
Functional morphology of the forelimb ...
Functional morphology of the pectoral girdle .. .
Comparative anatomy of the forelimb ...
Hindlimb : forelimb ratios ...
The evidence from preserved track ways ...
Summary : the probable function of the forelimb of lguanodon ...
Growth : the limbs of lguanodon bemissartensis
Individual variation and sexual dimorphism
E. - SUMMARY
F. - REFERENCES
G. - INDEX OF ABREVIATIONS .
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INST ITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE
KONINKLIJK BELGISCH INSTITUUT VOOR NATUURWETENSCHAPPEN
