1. Evolutionary Biology

Spinosaurus is not an aquatic dinosaur

  1. Paul C Sereno  Is a corresponding author
  2. Nathan Myhrvold
  3. Donald M Henderson
  4. Frank E Fish
  5. Daniel Vidal
  6. Stephanie L Baumgart
  7. Tyler M Keillor
  8. Kiersten K Formoso
  9. Lauren L Conroy
  1. 1Department of Organismal Biology, University of Chicago, United States
  2. Committee on Evolutionary Biology, University of Chicago, United States
  3. Intellectual Ventures, United States
  4. Royal Tyrrell Museum of Palaeontology, Canada
  5. Department of Biology, West Chester University, United States
  6. Grupo de Biología Evolutiva, UNED, Spain
  7. Department of Earth Sciences, University of Southern California, United States
  8. Dinosaur Institute, Natural History Museum of Los Angeles County, United States
https://doi.org/10.7554/eLife.80092
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Cite this article
  1. Paul C Sereno
  2. Nathan Myhrvold
  3. Donald M Henderson
  4. Frank E Fish
  5. Daniel Vidal
  6. Stephanie L Baumgart
  7. Tyler M Keillor
  8. Kiersten K Formoso
  9. Lauren L Conroy
(2022)
Spinosaurus is not an aquatic dinosaur
eLife 11:e80092.
https://doi.org/10.7554/eLife.80092
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14 figures, 14 tables and 3 additional files

Figures

Figure 1
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Digital skeletal reconstructions of the African spinosaurids Spinosaurus aegyptiacus and Suchomimus tenerensis.

(A) S. aegyptiacus (early Late Cretaceous, Cenomanian, ca. 95 Ma) showing known bones based on the holotype (BSPG 1912 VIII 19, red), neotype (FSAC-KK 11888, blue), and referred specimens (yellow) and the center of mass (red cross) of the flesh model in bipedal stance (overlap priority: neotype, holotype, referred bones). (B) Cervical 9 (BSPG 2011 I 115) in lateral view and coronal cross-section showing internal air space. (C) Caudal 1 centrum (FSAC-KK 11888) in anterolateral view and coronal CT cross-section. (D) Right manual phalanx I-1 (UCRC PV8) in dorsal, lateral, and sagittal CT cross-sectional views. (E) Pedal phalanges IV-4, IV-ungual (FSAC-KK 11888) in dorsal, lateral, and sagittal CT. (F) S. tenerensis (mid Cretaceous, Aptian-Albian, ca. 110 Ma) showing known bones based on the holotype (MNBH GAD500, red), a partial skeleton (MNBH GAD70, blue), and other referred specimens (yellow) (overlap priority: holotype, MNBH GAD70, referred bones). (G) Dorsal 3 in lateral view (MNBH GAD70). (H) Left manual phalanx I-1 (MNBH GAD503) in dorsal, lateral, and sagittal CT cross-sectional views. (I) Caudal 1 vertebra in lateral view (MNBH GAD71). (J) Caudal ~3 vertebra in lateral view (MNBH GAD85). (K) Caudal ~13 vertebra in lateral view with CT cross-sections (coronal, horizontal) of the hollow centrum and neural spine (MNBH GAD70). ag, attachment groove; C2, 7, 9, cervical vertebra 2, 7, 9; CA1, 10, 20, 30, 40, caudal vertebra 1, 10, 20, 30, 40; clp, collateral ligament pit; D4, 13, dorsal vertebra 4, 13; dip, dorsal intercondylar process; k, keel; mc, medullary cavity; nc, neural canal; ns, neural spine; pc, pneumatic cavity; pl, pleurocoel; r, ridge; S1, 5, sacral vertebra 1, 5. Dashed lines indicate contour of missing bone, arrows indicate plane of CT-sectional views, and scale bars equal 1 m (A, F), 5 cm (B, C), 3 cm (D, E, H–K) with human skeletons 1.8 m tall (A, F).

Figure 2
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Digital flesh model of Spinosaurus aegyptiacus.

(A) Translucent flesh model in hybrid swimming pose showing centers of mass (red cross) and buoyancy (white diamond). (B) Opaque flesh model in axial swimming pose with adducted limbs. (C) Modeled air spaces (‘medium’ option) include pharynx-trachea, lungs and paraxial air sacs. (D) Wading-strike pose at the point of flotation (2.6 m water depth) showing center of mass (red cross) and buoyancy (white diamond). lu, lungs; pas, paraxial air sacs; tr, trachea.

Figure 3
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Biomechanical evaluation of Spinosaurus aegyptiacus in water.

(A) Tail thrust (yellow curve) and opposing drag forces as a function of swimming velocity at the surface (blue) and submerged (green), with drag during undulation estimated at three and five times stationary drag. (B) Stability curve for the flesh model of S. aegyptiacus in water showing torque between the centers of mass (red cross) and buoyancy (white diamond), unstable equilibria when upright or upside down (positions 1, 5), and a stable equilibrium on its side (position 3) irrespective of the volume of internal air space. Curves are shown for flesh models with minimum (magenta) and maximum (green) air spaces with a dashed line showing the vertical body axis and vector arrows for buoyancy (up) and center of mass (down).

Figure 4
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Skeletal comparisons between Spinosaurus aegyptiacus, a basilisk lizard and secondarily aquatic vertebrates.

(A) Tail in S. aegyptiacus showing overlap of individual neural spines (red) with more posterior vertebral segments. (B) Sail structure in the green basilisk (CT-scan enlargement) and in vivo form and coloration of the median head crest and sail (Basiliscus plumifrons FMNH 112993). (C) Structure of the tail fluke in a urodele, mosasaur, crocodilian, and whale. (D) Centrum proportions along the tail in the northern crested newt (Triturus cristatus FMNH 48926), semiaquatic lizards (marine iguana Amblyrhynchus cristatus UF 41558, common basilisk Basiliscus basiliscus UMMZ 121461, Australian water dragon Intellagama lesueurii FMNH 57512, sailfin lizard Hydrosaurus amboinensis KU 314941), an extinct mosasaurid (Mosasaurus sp. UCMP 61221; Lindgren et al., 2013), an alligator (Alligator mississippiensis UF 21461), and Spinosaurus (S. aegyptiacus FSAC-KK 11888). Data in Appendix 2.

Figure 5
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Appendage versus total body surface area in aquatic and semiaquatic vertebrates.

Spinosaurus aegyptiacus and other non-avian theropods (green polygon, centroid large diamond) have appendages with considerable surface area compared to aquatic and semiaquatic vertebrates (blue polygon, centroid large dot). Underwater fliers (1–7 circled), which propel themselves with lift-based wings, also have less overall appendage surface area than in S. aegyptiacus and other non-avian theropods. Underwater fliers: 1, plesiosaur Cryptoclidus oxoniensis; 2, leatherback sea turtle Dermochelys coriacea; 3, emperor penguin Aptenodytes forsteri; 4, sea lion Zalophus californianus; 5, elasmosaur Albertonectes vanderveldei; 6, nothosaur Ceresiosaurus calcagnii; 7, pliosaur Liopleurodon ferox.

Figure 6
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Appendage surface area and scaling of paddle surface areas in crocodylians compared to S. aegyptiacus.

(A) Right hind foot of Spinosaurus aegyptiacus (FSAC-KK 11888) showing the outlines of digital flesh based on the living ostrich (Struthio camelus) as well as partial (pink) and full (blue) interdigital webbing. (B) Hind foot of an adult Alligator mississippiensis (WDC) in ventral view. (C) Forefoot of an adult A. mississippiensis (WDC) in ventral view. (D) Tail of an adult A. mississippiensis (WDC) in lateral view with CT visualization of vertebrae within the fleshy tail fluke. (E) Log-log plot of surface areas of webbed hind foot and side of the tail as a function of total body area in a growth series for A. mississippiensis (hind foot, green dots; tail, blue diamonds) and adult S. aegyptiacus (hind foot, purple-blue dots; tail, yellow diamond). I, IV, V, digits I, IV, V; un, ungual. Scale bars are 10 cm (A) and 3 cm (B–D).

Figure 7
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Paleogeographic location of spinosaurid fossils.

(A) Paleogeographic map (early Albian, ~110 Mya; Scotese, 2014). showing the circum-Tethyan fossil localities for baryonychines (Baryonyx, Suchomimus) and spinosaurines (Ichthyovenator, Vallibonavenatrix, Oxalaia, Irritator/Angaturama, Spinosaurus). Spinosaurus localities (yellow asterisks) range across northern Africa from coastal (sites 1, 2) to inland (site 3) sites. (B) Spinosaurus sp. right maxilla (MNBH EGA1) from Égaro North (central Niger) in medial (top) and ventral (bottom) views and shown (red) superposed on the snout of Spinosaurus aegyptiacus. 1, S. aegyptiacus holotype (Bahariya, Egypt); 2, S. aegyptiacus neotype (Zrigat, Morocco); 3, Spinosaurus sp. (Égaro North, Niger); am, articular rugosities for opposing maxilla; aofe, antorbital fenestra; Ba, Baryonyx walkeri; en, external naris; Ic, Ichthyovenator laosensis; Ir, Irritator challengeri/Angaturama limai; m3, 12, maxillary alveolus 3, 12; Ox, Oxalaia quilombensis; Su, Suchomimus tenerensis; t, tooth; Va, Vallibonavenatrix cani. Scale bar is 10 cm.

Figure 8
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Calibrated phylogeny of spinosaurids (Barremian to Cenomanian, ~35 My).

Updated phylogenetic analysis of spinosaurids resolves two stages in the evolution of piscivory and display. We show key cranial adaptations in the skull and highlight changes at the anterior end of the trunk to enhance neck ventroflexion (second dorsal vertebra in lateral and anterior views). Bottom, the fully terrestrial theropod Allosaurus fragilis (Madsen, 1976); middle, the baryonychine spinosaurid Suchomimus tenerensis (MNBH GAD70); top, the spinosaurine Spinosaurus aegyptiacus (BSPG 1912 VIII 19). con, condyle; dr, dental rosette; en, external naris; hy, hypopophysis; k, keel; ncr, nasal crest; ns, neural spine; pa, parapophysis; tp, transverse process.

Figure 9
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Comparison of skeletal reconstructions for Spinosaurus aegyptiacus in left lateral view.

(A) Digital skeletal reconstruction from this study in left lateral view. (B) Pelvic girdle. (C) Cervical column (C1–10). (D) Pectoral girdle and forelimb. (E) Hind limb. (F) Anterior trunk. (G) Silhouette skeletal drawing from the aquatic hypothesis (from Ibrahim et al., 2020b). On one side of each length comparison, one or two blue lines are shown that register the alternative reconstructions. The opposing end of each length comparison either has a single blue line (when comparisons match, both 100%) or a red line as well for the shorter one (<100%): A blue line on the right or top sides of each comparison is used for registration. The opposing side has a blue line if reconstructions agree on length (100%), or a blue line for the length estimate in this study and a red line for that of the aquatic hypothesis. In all disparate comparisons, the reconstruction in this study is shorter (percentage given). Skeletal reconstructions (A, G) are aligned by the anterior and posterior margins of the ilium and measured to the cervicodorsal junction (C10-D1); the pelvic girdle (B) is aligned along the ventral edge of the sacral centra and base of the neural spines and measured to the distal ends of the pubis and ischium; the cervical column (C) is aligned at the cervicodorsal junction (C10-D1) and measured to the anterior end of the axis (C2); the scapula and components of the forelimb (humerus, ulna, manual digit II, manual phalanx II-1) (D) are aligned at the distal end of the blade and their proximal ends, respectively, and measured to the opposing end of the bone; the components of the hind limb (femur, tibia, pedal digits I, III) (E) are aligned at their proximal ends and measured to the opposing end of the bone; and anterior trunk depth (F) is aligned along the ventral edge of the centrum and neck of the spine of D6 and measured to the ventral edge of the coracoid. II-1, manual phalanx II-1; Ili, ilium; F, femur; H, humerus; Ish, ischium; l, left; Pe, pes; Pu, pubis; Ma, manus; r, right; RaU, radius-ulna; Sc, scapula; TF, tibia-fibula.

Figure 10
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CT scans inform cross-sectional muscle mass in Spinosaurus aegyptiacus.

Muscle mass reconstructions of the axial column at five points (A-E) in S. aegyptiacus are compared to CT scan cross-sections of Struthio (Snively and Russell, 2007) and Alligator (Wedel, 2003; Mallison et al., 2015).

Figure 11
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Cross-sections from a CT scan of Basiliscus plumifrons FMNH 112993.

(A) Skeleton showing position of CT sections of the axial column. (B) Anterior cervical region (C2). (C) Mid cervical region (C3). (D) Posterior cervical and anterior dorsal region (C4-D1). (E) Mid dorsal region (D12). (F) Anterior caudal region (CA4). (G) Caudal region at most posterior transverse process (CA10). (H) Mid caudal region (CA15). CT scan data available on Morphosource.org. Scale bars, 5 mm.

Appendix 4—figure 1
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Anterior dorsal centrum referable to Spinosaurus sp. (from Brusatte and Sereno, 2007: Fig. 9).

Photographs and line drawings of MNBH IGU11 from the Echkar Formation (Cenomanian) of Niger in ventral (A) dorsal (B), right lateral (C), and posterior (D) views. Cross-hatching indicates broken bone. ana, articular surface for the neural arch; hy, hypapophysis; k, ventral keel; nc, neural canal; pa, parapophysis; pc, pleurocoel. Scale bar, 5 cm.

Appendix 5—figure 1
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Phylogenetic tree for Spinosauroidea.

Single most-parsimonious tree for 12 spinosauroid terminal taxa (9 spinosaurids) and 120 characters split evenly between the cranium (49%) and postcranium (51%), showing decay values (Consistency index = 0.81, Retention Index = 0.85).

Appendix 5—figure 2
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Posterior skull roof of the baryonychine spinosaurid Suchomimus tenerensis (field no. GAD302).

Composite restoration of the posterior skull roof of Suchomimus tenerensis in (A) lateral and (B) dorsal views showing a swollen postorbital brow and narrow orbital notch limiting the frontal orbital margin. Based on original bone (blue), reflected original bone (gray), and reconstructed paroccipital processes.

Tables

Table 1
Relative size of specimens in the skeletal reconstruction of S. aegyptiacus.

Relative sizes of key specimens used in the skeletal model of S. aegyptiacus (nos. 1–4) and select bones (nos. 5, 6) from Egypt and Morocco. All are scaled to the size of the adult snout (MSMN V4047).

No.SpecimenMaturityRelative size (%)LinearupsizingDescription
1BSPG
1912 VIII 19		Subadult761.32Holotype (destroyed) preserving dentaries, presacral, sacral and caudal vertebrae including the dorsal sail (Stromer, 1915)
2FSAC-KK 11888Subadult761.32Neotype preserving skull bones, partial limbs, dorsal sail and most of the tail (Ibrahim et al., 2014)
3BSPG
1922X45		Subadult661.51‘Spinosaurus B’ (destroyed) fragmentary specimen with five partial dorsals (~D1 centrum, mid dorsal centrum, partial ~D13 vertebra), seven partial caudal vertebrae, and both tibiae (Stromer, 1934)
4MSMN V4047Adult100—Isolated snout with broken teeth (Dal Sasso et al., 2005); large size and coossified sutures indicate maturity
5UCRC PV8Adult801.25Large manual phalanx I-1 (28.0 cm length) of an adult within reach but still smaller than the 35 cm length estimated on the basis of the proportions in the manus of Angaturama (?=Irritator) scaled to the adult snout (Aureliano et al., 2018)
6UCRC PV24Adult751.33Large Kem Kem vertebra, ~C9, from Gara Sbaa (centrum length 11.6 cm, centrum width 14.0 cm)
Table 2
Axial muscle area in the crested basilisk.

Area measurements of epaxial and hypaxial musculature along the axial column in the crested basilisk Basiliscus plumifrons (FMNH 112993). C, cervical; D, dorsal; CA, caudal.

LocationTotal areaEpaxial areaHypaxial area
mm2mm2% totalmm2% total
Anterior neck (C2)180.483.946.596.553.5
Mid neck (C3)139.754.939.384.860.7
Posterior neck (C4-D1)202.169.834.7132.365.3
Mid dorsal (D12)212.060.028.3152.071.7
Basal tail (CA4)434.0153.835.4280.264.6
Anterior tail (CA10, last transverse process)221.763.528.6158.271.4
Mid tail (CA15)97.829.829.868.070.2
Table 3
Axial muscle and transverse process length in the tail of the crested basilisk.

Transverse processes versus muscle width in the tail cross-sections in the crested basilisk Basiliscus plumifrons (FMNH 112993). Measurements are from the midline to the distal end of the transverse process (or centrum margin when there is no process) and to the lateral surface of the tail. CA, caudal.

LocationMeasurementTotal(mm)% bone and muscle% muscle only
CA4Transverse process width16.862.937.1
Total width26.7
CA10Transverse process width8.15743.0
Total width14.2
CA15Centrum width3.536.863.2
Total width9.5
Table 4
Epaxial muscle height and neural spine height in the tail of the crested basilisk.

Height of neural spines versus epaxial musculature in tail cross-sections in the crested basilisk Basiliscus plumifrons (FMNH 112993). Measurements are from the dorsal surface of the centrum to the top of the epaxial muscle and to the distal end of the neural spine. CA, caudal.

LocationMeasurementTotal(mm)% neural spine adjacent to muscle% neural spine above muscle
CA4Neural spine height18.470.729.3
Epaxial mm height13.0
CA10Neural spine height33.632.167.9
Epaxial mm height10.8
CA15Neural spine height34.523.276.8
Epaxial mm height8.0
Table 5
Hypaxial muscle depth and chevron length in the tail of the crested basilisk.

Chevron length versus hypaxial muscle depth in tail cross-sections in the crested basilisk Basiliscus plumifrons (FMNH 112993). Measurements are from the ventral surface of the centrum to the distal tip of the chevron and to the ventral surface of the tail. CA, caudal.

LocationMeasurementTotal(mm)% chevron length% muscle below chevron
CA4Chevron depth9.052.647.4
Hypaxial depth17.1
CA10Chevron depth7.167.033.0
Hypaxial depth10.6
CA15Chevron depth4.867.632.4
Hypaxial depth7.1
Table 6
Density, volume, and mass in the flesh model of S. aegyptiacus.

Whole-body and body part densities, volumes, and masses for the new mesh adult flesh model of S. aegyptiacus.

No.Body partitionAverage density(kg/m3)% of axial volume without sailMass(kg)
1Whole body833—7390
2Axial body
(excluding lung/sail)		788100.05794
3Axial head-trunk
(not lung/sail/tail)		85064.83209
4Axial tail
(not sail)		100035.22585
5Forelimb (paired)10503.8108
6Hind limb (paired)105011.1590
7Dorsocaudal sail11968.5441
8Lungs012.50
Table 7
Flesh model functional dimensions in S. aegyptiacus.

Functional dimensions for the adult flesh model of S. aegyptiacus in sculling pose.

No.DimensionMeasure (m)
1Total body length (sculling pose)13.53
2Body length minus tail (sculling pose)6.92
3Head length1.57
4Neck length (sculling pose)2.18
5Trunk depth (mid trunk, without sail)1.28
6Trunk sail depth (mid trunk)1.93
7Trunk sail length (maximum at base)3.53
8Tail length6.61
9Tail depth at base1.38
10Tail depth at midpoint0.97
11Tail depth at distal end0.87
12Tail depth average1.08
13Forelimb length (straightened)1.85
14Hind limb length (straightened)2.88
Table 8
Flesh model volume and surface area in S. aegyptiacus.

Adult flesh model whole-body and body part volumes and surface areas as measured in MeshLab. Surface area of body parts does not include cut surfaces.

No.Body partVolume (m3)Surface area (m2)
1Whole body8.9454.06
2Body above waterline (floating)1.6522.58
3Body below waterline (floating)7.2731.38
4Head0.212.23
5Neck0.784.37
6Trunk4.0111.44
7Trunk sail (both sides, external edge)0.4010.06
8Forelimb (both)0.243.86
9Hind limb (both)0.455.29
10Tail2.8116.56
11Tail with axial muscle2.7113.27
12Tail sail only0.103.17
13Airspace-minimum
(~4% body volume)		0.374.86
14Airspace-medium
(~8% body volume)		0.676.63
15Airspace-maximum
(~12% body volume)		1.088.66
Table 9
Center of mass (CM) calculations for S. aegyptiacus.

There have been four estimates for the location of CM in flesh models of S. aegyptiacus using four different points of origin as a reference. Because three were based on an adult flesh model, we convert the one study based on a subadult (number 3) to reflect its position in an adult flesh model: “x” is the distance anterior tothe origin, and “y” is the height above the ground, both in cm.

No.Author/resultx-originxyNotes
1Ibrahim et al., 2014; quadrupedHip joint>81–Based on an adult flesh model, no coordinates given, CM shown graphically under D10 and said to be anterior to hip/knee joints at a distance greater than femur length (MeshLab calculation error)
2Henderson, 2018; bipedTip of tail8,850100Based on an 3D mesh model based on the adult skeletal model of Ibrahim et al., 2014 with estimated length 16 m long; y-coordinate origin is ‘lowest point of axial body’
3Ibrahim et al., 2020b; quadruped‘Cranial rim’ of acetabulum72.5–82.5
(adult = 95.7–108.9)		–81
(adult: 106.9)		Based on a flesh model of the subadult neotype with femur length of 62.5 cm (actual 61.0 cm); y-coordinate measures to substrate
4This paper; bipedApex of the acetabulum15.3–240Based on an adult flesh model with avian-style internal air spaces and femur length of 81.0 cm; y-coordinate measures to substrate
Table 10
Estimated internal air space in S. aegyptiacus.

Air space options for the adult flesh model of S. aegyptiacus and their effect on whole-body density, body mass (BM), and center of mass (CM). The x-coordinate for CM is measured from the apex of the acetabulum.

No.Air space optionPart of whole-body volume (%)Mean whole-body density(g/l)BM(kg)CM x-coordinate(cm)
1Minimum
(lizard-like)		4.0909801328.5
2Medium
(croc-like)		8.0875771623.2
3Maximum
(bird-like)		12.5833739015.3
Appendix 2—table 1
Extant newt and squamate specimens used for muscle mass estimation and for logging centrum proportions along the tail.

Specimen numbers have hyperlinks to MorphoSource specimen pages with associated scans and 3D models.

Specimen no.TaxonSexTypePurpose
FMNH 84926Triturus cristatusMaleWetCaudal centrum ratios, muscle mass estimation
FMNH 57512Intellagama lesueuriiMaleWetMuscle mass estimation
FMNH 22389Intellagama lesueuriiMaleSkeletonCaudal centrum ratios, muscle mass estimation
KU 314941Hydrosaurus amboinensisUnknown; low neural spines (female/ immature male)WetCaudal centrum ratios
FMNH 52698Hydrosaurus pustulatusMaleWetMuscle mass estimation
FMNH 236131Hydrosaurus pustulatusUnknown; low neural spines (female/ immature male)SkeletonMuscle mass estimation
FMNH 112993Basiliscus plumifronsMaleWetMuscle mass estimation
FMNH 257162Basiliscus plumifronsMaleSkeletonAdditional high-resolution visualization
UMMZ 121461Basiliscus basiliscusMaleWetCaudal centrum ratios
UF 41558Amblyrhynchus cristatusUnknownWetCaudal centrum ratios
FMNH 22042Amblyrhynchus cristatusUnknown;?male (tallest caudal neural spines in FMNH collection)SkeletonMuscle mass estimation
UF 21461Alligator mississippiensisUnknown; juvenileWetX-ray image of caudal centra in tail; caudal centrum ratios
Appendix 2—table 2
Caudal centrum ratios along the tail in S. aegyptiacus, Mosasaurus, and extant semiaquatic amphibians and reptiles (Figure 4D).
L-H ratiosRelative distance (caudal #/total # of caudals)
CA #Spino.Mosa.Ambly.Alli.Basil.Hydro.Trit.Intel.Spino.Mosa.Ambly.Alli.Basil.Hydro.Trit.Intel.
10.731.401.141.521.253.261.600.020.010.020.020.020.020.030.02
21.561.111.921.503.782.210.040.030.040.050.040.040.050.04
31.551.052.051.973.182.480.060.040.060.070.060.050.080.06
40.841.881.182.302.053.202.730.080.050.090.100.080.070.110.08
51.861.222.442.142.882.780.100.070.110.120.100.090.140.10
61.961.332.502.143.082.830.120.080.130.150.120.110.160.13
71.121.062.061.382.552.263.052.980.140.090.150.170.140.130.190.15
81.151.032.101.432.722.173.112.990.160.110.170.200.160.140.220.17
90.952.131.492.872.413.073.220.180.120.190.220.180.160.240.19
100.972.191.522.892.452.903.260.200.140.210.240.200.180.270.21
110.982.291.533.142.342.953.170.220.150.230.270.220.200.300.23
120.932.371.672.942.503.173.280.240.160.260.290.240.210.320.25
131.241.072.571.623.272.482.733.340.260.180.280.320.250.230.350.27
141.211.042.521.653.352.432.773.210.280.190.300.340.270.250.380.29
151.062.591.743.492.642.853.510.300.200.320.370.290.270.410.31
160.912.671.723.462.522.953.640.320.220.340.390.310.290.430.33
171.430.922.741.913.552.942.653.520.340.230.360.410.330.300.460.35
181.330.952.801.813.662.823.293.730.360.240.380.440.350.320.490.38
190.992.911.893.733.033.143.610.380.260.400.460.370.340.510.40
201.370.982.771.973.892.852.703.500.400.270.430.490.390.360.540.42
210.970.992.941.893.912.892.823.730.420.280.450.510.410.380.570.44
221.400.982.991.953.992.873.163.690.440.300.470.540.430.390.590.46
231.381.022.992.124.012.823.273.930.460.310.490.560.450.410.620.48
241.320.923.002.213.742.962.923.900.480.320.510.590.470.430.650.50
251.410.883.052.254.403.293.223.960.500.340.530.610.490.450.680.52
261.360.903.052.494.143.123.633.720.520.350.550.630.510.460.700.54
271.350.933.022.634.253.313.453.990.540.360.570.660.530.480.730.56
281.150.903.242.793.693.133.364.030.560.380.600.680.550.500.760.58
291.440.833.272.704.373.253.034.140.580.390.620.710.570.520.780.60
301.290.893.312.993.973.183.114.060.600.410.640.730.590.540.810.63
311.340.973.223.103.823.373.233.770.620.420.660.760.610.550.840.65
321.420.823.543.214.563.483.274.220.640.430.680.780.630.570.860.67
331.360.943.443.324.763.462.193.860.660.450.700.800.650.590.890.69
341.450.883.313.454.873.481.754.250.680.460.720.830.670.610.920.71
350.893.073.925.093.591.894.110.700.470.740.850.690.630.950.73
360.893.253.725.193.682.704.000.720.490.770.880.710.640.970.75
371.960.823.503.905.543.711.984.210.740.500.790.900.730.661.000.77
380.823.312.965.503.654.170.760.510.810.930.750.680.79
390.823.223.195.943.824.130.780.530.830.950.760.700.81
400.823.193.545.933.713.950.800.540.850.980.780.710.83
411.740.773.073.065.953.914.450.820.550.871.000.800.730.85
420.793.255.994.064.310.840.570.890.820.750.88
430.723.426.223.994.670.860.580.910.840.770.90
440.753.296.734.324.800.880.590.940.860.790.92
450.743.166.644.244.590.900.610.960.880.800.94
460.802.636.004.334.560.920.620.980.900.820.96
470.832.535.824.425.310.940.641.000.920.840.98
480.906.614.663.270.960.650.940.861.00
490.886.164.610.980.660.960.88
500.986.494.391.000.680.980.89
510.955.494.290.691.000.91
520.834.390.700.93
530.914.520.720.95
540.914.530.730.96
550.884.180.740.98
560.963.970.761.00
570.890.77
581.040.78
590.960.80
600.940.81
611.080.82
620.990.84
630.860.85
640.880.86
651.060.88
661.030.89
670.960.91
681.020.92
691.010.93
700.990.95
711.000.96
721.010.97
731.010.99
740.981.00
Appendix 3—table 1
Forefoot, hind foot, and tail area and other data in five species of extant crocodylians.

American alligator (1–7, Alligator mississippiensis); Schneider’s dwarf crocodile (8, Paleosuchus trigonatus); broad-snouted caiman (9, Caiman latirostris); spectacled caiman (10, Caiman crocodilus); African dwarf crocodile (11, Osteolaemus tetraspis).

No.Total length (cm)Snout-vent length(cm)Skull length(cm)Post skull width (cm)Tail length(cm)Mass (kg)Sex(m, f)Age est.(year)Hand(cm)Foot(cm)Tail(cm)
165.032.09.05.535.51.15f25.79.890.9
269.036.09.85.635.01.05f127.013.2102.8
385.042.511.56.843.01.95f310.923.6114.1
4105.452.214.08.554.53.65f418.339.9267.7
5154.078.022.014.377.515.45m1246.293.3655.2
6167.085.021.815.586.822.95m?51.496.0717.9
7211.5104.525.318.5108.039.10m2369.5146.51372.5
886.347.612.68.537.02.85f58.419.8172.1
975.538.08.66.338.01.55f35.818.3141.6
10108.855.514.010.055.54.95f512.129.6272.1
1168.233.39.46.232.41.15f46.317.199.0
Appendix 4—table 1
Fossil material referable to Spinosaurus from inland basins in Algeria and Niger. D, dorsal.
SpecimensDescription
MNHN SAM 124Snout with preserved length of 62 cm (Gara Samani, Béchar Basin, Algeria) compared to ~1 m for adult snout MSNM V4047
MNHN SAM 125Premaxilla fragment (Gara Samani, Béchar Basin, Algeria)
MNHN SAM 125–8Two cervical vertebra, one dorsal vertebra (Gara Samani, Béchar Basin, Algeria)
MNBH IGU11~D1 short, very low oval centrum with low parapophysis, very strong ventral keel (In Abangharit, Iullumeden Basin, Niger)
MNBH EGA1Both maxillae and a portion of the alveolar edge of the right dentary with tooth roots within alveoli (Égaro North, Chad Basin, Niger)
MNBH EGA2Isolated tooth with root (Égaro North, Chad Basin, Niger)

Additional files

MDAR checklist
https://cdn.elifesciences.org/articles/80092/elife-80092-mdarchecklist1-v1.pdf
Source data 1

Data matrix as nex file for Mesquite.

https://cdn.elifesciences.org/articles/80092/elife-80092-data1-v1.zip
Source data 2

Data matrix as tnt file for TNT.

https://cdn.elifesciences.org/articles/80092/elife-80092-data2-v1.zip

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  1. Paul C Sereno
  2. Nathan Myhrvold
  3. Donald M Henderson
  4. Frank E Fish
  5. Daniel Vidal
  6. Stephanie L Baumgart
  7. Tyler M Keillor
  8. Kiersten K Formoso
  9. Lauren L Conroy
(2022)
Spinosaurus is not an aquatic dinosaur
eLife 11:e80092.
https://doi.org/10.7554/eLife.80092